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Multivariate analysis of the Carex brevior group in Iowa

Scott C. Zager University of Northern Iowa

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MULTIVARIATE ANALYSIS OF THE CAREX BREVIOR GROUP

IN IOWA

An Abstract of a Thesis

Submitted

In Partial Fulfillment

of the Requirements for the Degree

Master of Arts

Scott C. Zager

University of Northern Iowa

December 1991

LI BRARY UNIVERSITY OF NORTHERN IOWA CEDAR FALLS, IOWA ABSTRACf

In Iowa, the most troublesome sedges to identify are Carex brevior, .,C. festucacea, ..C. molesta, ..C. normalis, ..C. tenera, and ..C. tenera var. echinodes.

These taxa form the _c. brevior group--part of an even larger aggregate of species associated with _c. straminea. Their morphological features are indistinct and intergrade into one another. Botanists have had difficulty classifying members of this aggregate for nearly 200 years. Some authors have viewed the taxa as separate species, others as polymorphic forms of a single species. Much of the contention has centered upon the variation of taxonomic characters used to delimit and distinguish species.

The objectives of my study were: 1) to randomly sample up to 30 specimens from each population, at several populations of each taxa, to obtain statistical parameters for morphological characters and ascertain if variation occurs within populations, between populations, and between taxa; 2) to evaluate

44 characters and 12 character ratios for each taxa by Univariate Analysis and

Stepwise Discriminant Analysis (SDA) and obtain reliable taxonomic characters or character combinations; and 3) to test the validity of taxonomic classifications within the _c. brevior group by determining if the morphological forms are significantly different using Canonical Discriminant Analysis (CDA).

There were 450 samples collected at 15 sites (21 populations). The taxa were mostly found in different micro-habitats. Most character means were significantly different (l! < 0.0001), e.g., means of _c. molesta and _c. brevior were significantly different for 52 of the 56 characters tested. However, single characters could not reliably separate taxa because of overlapping ranges of variation. CDA revealed taxa to be significantly different (F = 24.08; ~ <

0.0001) along 4 canonical axes. SDA identified character combinations or suites which could reliably delimit and distinguish taxa. There were no subgroups observed within the taxa. Most of the variation expressed by each taxon was found within populations and there were few differences between populations.

The characters varied in predictable patterns and this variation is mostly attributed to phenotypic plasticity. However, specimens were found at sympatric sites with unusual character states or mixed character suites, suggesting they are putative hybrids. .,:)..2 T - I '1

MULTIVARIATE ANALYSIS OF THE CAREX BREVIOR GROUP

IN IOWA

A Thesis

Submitted

In Partial Fulfillment

of the Requirements for the Degree

Master of Arts

Scott C. _?,ager

University of Northern Iowa

December 1991

LIBRARY UNIVERSITY OF NOf:THE RN IOWA CEDAR FALLS, IOWA ii

This Study by: Scott C. Zager

Entitled: MULTIVARIATE ANALYSIS OF THE CAREX BREVIOR GROUP

IN IOWA

has been approved as meeting the thesis requirement for the

Degree of Master of Arts.

~~"'I Date Dr. Lawrence J. Eilers, Chair, Thesis Committee '1/.zsJ; Date Dr. Daryl D. Smith, Member, Thesis Committee

Y/10! ~ Date Dr. JaneL. Kotenko, Member, Thesis Committee

1/ti~( Date

(2 -20-(l Date Dr. John W. Somervill, Dean, Graduate College

·· -\~~ "··-- -- , 1,..1., ~ t.lV f::,: ~ '.,.' . ~. : i~:. 1, ,'""'t ~" ( '""' 'lf~r ~·rr~):1Y {~F r· :C~t~T:·~~·f~ 1 ·~~ 10\~]i i r:r:i:.t;:. f\~ F;'.Li.::;. tf)V~·' :j lll

ACKNOWLEDGEMENTS

I have the privilege of being Dr. Lawrence J. Eilers' last graduate student; an honor he bestowed upon me for two years after his retirement. Mere words cannot express the magnitude of my gratitude nor my appreciation for his allowing me to perform this study under his direction. While primarily a floristic botanist, he has supervised my study with great insight, always focusing my thoughts toward the principles of good scientific research. He has been a true mentor in every sense. If this study has any significance, it is because he has

disciplined my thoughts. If it has any error, it is because he would not restrain

my enthusiasm. He has been a leader in preserving Iowa's diminishing natural

areas, but he has expressed that his most cherished legacy is the students he has

encouraged to become botanists, ecologists and conservationists. I was also

fortunate to have assistance from Dr. Kay Klier, a plant taxonomist, whose career

I predict will be equally as rewarding to the students she instructs.

I wish to express my gratitude to my committee members Dr. Jane

Kotenko and Dr. Paul Christiansen. I especially wish to thank Dr. Daryl Smith

who gave me the opportunity to study at the University of Northern Iowa. All

are individuals which I greatly admire.

I wish to thank other University of Northern Iowa faculty: Dr. P. Whitson,

Dr. 0. Schwartz, Dr. A Orr and Dr. J. Trainer.

I wish to acknowledge The Iowa Academy of Science for funding travel

expenses during this study. I also wish to thank Donna Ware and the Delzie

Demaree Travel Award Committee for the opportunity to visit the Missouri iv

Botanical Gardens. I wish to thank the College of Natural Sciences, University of

Northern Iowa, for the Lantz Fellowship Award which enabled me to initiate this study.

I wish to thank all the curators for loans, especially Dr. P. Holmgren (NY) and A E. Schuyler (PH) for their hospitality. Also, I wish to thank Dr. C.

Reimers for his assistance in Philadelphia. I gratefully acknowledge Dr. P.

Rothrock for his correspondence and assistance in identifying important taxa. I deeply appreciate Dr. A A Reznicek for his clarification of Carex straminea.

Other individuals I wish to thank include Mark Leoschke, Barbara Wilson and

John Nehnevaj for field assistance. I wish to thank AI Ehley, and the Roadside

Assistance Office for a place to write.

I wish to thank the Iowa Department of Natural Resources for permission

to collect in State Parks and the Iowa State Preserves Advisory Board for

permission to collect on Iowa preserves. More thanks to The Iowa Nature

Conservancy for reviewing my study and permission to visit their preserves, and

the Iowa-Illinois Gas and Electric Company for permission to visit Big Sand

Mound Preserve.

Finally, I would like to express my special appreciation to my wife Patty

Vander Waal, without whose patience, love, and sacrifice this study would not

have been possible. I wish also to thank my parents for their encouragement and

support, especially my father who recognized that the world needs good biologists. v

TABLE OF CONTENTS ACKNOWLEDGEMENTS . . . i i i

LIST OF TABLES . . . X LIST OF FIGURES ... xii CHAPTER

I. INTRODUCTION ...... 1 Statement of Problem .. 2 Taxonomic Classifications. 6 The Genus Carex ...... 6

Morpho 1ogy . . 6 Subgeneric classifications .... 7

The Subgenera of Carex . . 9 The Section Ovales Kunth ...... 10 The Subsection Festucaceae Mackenzie (nomina nuda}...... 11 The Carex brevior Group...... 11 The Taxonomic History of the Carex brevior Group . . . 13 Taxonomy of Carex straminea...... 14 Taxonomy of Carex brevior...... 16 Taxonomy of Carex tenera . 17 Taxonomy of Carex tenera var. echinodes. 18 Taxonomy of Carex festucacea . . . . 18 Taxonomy of Carex normalis .... 19 Taxonomy of Carex molesta 20 Carex Evolution, Morphology and Genetics ...... 20 Phylogenetic Development of the Inflorescence. . 20 vi

Phenotypic Plasticity and Meristematic Development ... 24

Genetics ...... 26

Cytotogenetics ... 26

Agmatoploidy. 26

Hybridization ... 28 Population genetics of species aggregates. 30 Genetics of the Carex brevior group. . . . 31 Summary of Literature...... 32 CHAPTER 2. METHODS ...... 33 Herbarium Study. 33

Field Collections ...... 34 Laboratory Evaluation .... . 36 Statistical Analyses .... . 36 Univariate Analyses .. . 36 Multivariate Analyses ... . 40 CHAPTER 3. RESULTS ...... 45 Field Observations .. 45 Growth Habit of Iowa Ovales 45

Ecological Characteristics . 49

Field observations of ~. brevior .. 50

Field observations of ~. molesta . 50

Field observations of ~. festucacea ... 52

Field observations of ~. tenera...... 52 vii

Field observations of~. normalis •. 53

Field observations of ~. tenera var. echinodes . 54 Statistical Analyses ...... 55 Univariate Analyses ...... 55 Test of normality. 55 Infraspecific variation .. 78 Univariate tests of taxonomic significance 78 Multivariate Analyses. 79 Multivariate tests of taxonomic significance . 79 Infraspecific variation .. 81 Taxonomic Characters ..... 81 Stepwise Discriminant Analysis 89 Morphological Descriptions ... 91 Carex molesta .. 92 Carex brevior ...... 93 Carex normalis ...... 96 Carex festucacea . . . . . 96 Carex tenera 96 Carex tenera var. echinodes. 97 Intermediate Specimens With Mixed Character Suites .... 97 Intermediate Specimens of ~. brevior and ~. molesta .. 99 Intermediate Specimens of ~. molesta and ~. festucacea 100 Intermediate Specimens of ~. brevior and ~. festucacea 101 ~--- ..

viii

Intermediate Specimens of ~. tenera and~. normalis. 102 CHAPTER 4. DISCUSSION • • ...... 103 Taxonomic Classifications...... 103 Taxonomic Characters .... 105 Vegetative Characteristics ... 105 Variation of vegetative characters . . . . 107 Inflorescence Characteristics ..... 108 Variation of the inflorescence 109 Spike Characteristics. 109 Variation of spike characters .. . 110 Scale and Bract Characteristics ... . 111 Variation of scale and bract characters. . 112 Perigynium Characteristics ...... 112 Characteristics of the Achene...... 114 Sources of Morphological Variation .... 115 Phenotypic Plasticity ... 115 Hybidization ...... 116 LITERATURE CITED 119 APPENDIX

A. Nomenclator of the ~. brevior group. 127

~. straminea ..... 128

~. festucacea .. . 129

~. tenera .. . 130

~. tenera var. echinodes .. 132 !7£1 ' 'elSalOW 'J

££1 • s ~ L'eW.AOU • J

Z£1 ' .AO~Aa..tq 'J

X~ ------

X

LIST OF TABLES Table

1. The Iowa taxa of the Carex brevior group (Cyperaceae) . 2

2. Taxa of the Carex, section Ovales in Iowa (Gilly 1946). 10 3. Suite of characters used in scoring specimens for analysis. Vegetative characters were excluded from multivariate analysis. Ratios were derived from measured attributes of characters for each specimen (see Figures 1, 2, and 3) ...... 37 4. Identification codes, site names and county location. 46 5. Taxon, site code, canonical plot site code, sample size, other species prescence (mixed or sympatric populations), and date collected ...... 47

6. Character description, taxon, sample size (N), mean, standard deviation (SO), minimum value and maximum value...... 56 7. Eigenvalue and the percentage of the total variation explained by each canonical axis ...... 89 8. Stepwise discriminant characters separating Carex brevior, £. festucacea, £. molesta, £. normalis, £. tenera, and £. tenera var. echinodes. 90 9. Stepwise discriminant characters separating the Carex brevior group into two subgroups for dichotimous key construction: Subgroup A = £. brevior and £. molesta; Subgroup B = £. festucacea, £. normalis, £. tenera and £. tenera var. echinoides ..... 92

10. Stepwise discriminant characters separating Carex festucacea, £. normalis, £. tenera and£. tenera var. echinoides into two pairs: Pair 1 = £. normalis and£. tenera var. echinoides; Pair 2 = £. festucacea and £. tenera ..... 93 11. Stepwise discriminant characters for separating Carex brevior and £. molesta ...... 94 12. Stepwise discriminant characters for separating Carex festucacea and £. tenera ...... 94 xi

13 . Stepwise disciminant characters for separating Carex brevior and £. festucacea ...... 95 14. Stepwise discriminate characters for separating Carex normalis and£. tenera var. echinoides . . . . . 95 15. Specimens displaying mixed character suites as identified by (1) mixed classifications of SAS DISCRIM procedure; (2) intermediate location between taxon classes on canonical plots; and (3) observation of atypical character combinations ...... 98 xii

LIST OF FIGURES Figure 1. Symmetric plane diagrams adapted from Radford et al. (1974). Horizontal numbers correspond to character states given in Table 3: numbers 40 and 53. Columnar numbers correspond to character states given in Table 3: 42 and 54...... 40 2. Perigynia of the Carex brevior group (redrawn from Mackenzie (1940) except for£. tenera which is redrawn from Boott (1862)). (a) Dorsal surface of perigynium, (b) ventral surface of perigynium, subtending perigynium scale, and (d) achene...... 41 3. Features of the inflorescence and spike (adapted from Mackenzie (1940)) ...... 42 4. Aerial views of rhizome growth patterns depicting locations of meristems and ramets along rhizome axes. (A) Initial culm developed from seed. (B) Rhizome elongating in 2 directions with 2 apical meristems and 6 culms. (C) Rhizome elongating in one direction with 1 apical meristem and 15 culms. (D) Rhizome appendages elongating in several directions at ends of 4 branches (rhizome has many meristems and culms at the ends, but is barren at center). (E) Fragmented rhizome branches elongating in one to many directions away from the origin...... 48 5. Box plots of selected characters of Carex brevior (B), £. tenera var. echinodes (E), £. festucacea (F), £. molesta (M), £. normalis (N), and£. tenera. Means indicated by heavy line; mode by capped line; box indicates interquartile (25th-75th percentile). . . . . 70

6. Plot of canonical axes 1 and 2 showing ordination of Carex brevior (B), Carex festucacea (F), Carex molesta (M), Carex normalis (N), Carex tenera (T), and Carex tenera var. echinodes (E). Taxon clusters were significantly different at F = 24.08. . . 82

7. Plot of canonical axes 2 and 3 showing ordination of Carex brevior (B), Carex festucacea (F), Carex molesta (M), Carex normalis (N), Carex tenera (T), and Carex tenera var. echinodes (E). Taxon clusters were significantly different at F = 24.08. . . 83 Xi i i

8. Plot of canonical axes 3 and 4 showing ordination of Carex brevior (B), Carex festucacea (F), Carex molesta (M), Carex normalis (N), Carex tenera (T), and Carex tenera var. echinodes (E). Taxon clusters were significantly different at F = 24.08. . . 84 9. Plot of canonical axes 1 and 2 showing ordination of specimens according to population codes of collection sites (Table 5) (Compare with Figure 6). . . 85 10. Plot of canonical axes 2 and 3 showing ordination of specimens according to population codes of collection sites (Table 5) (Compare with Figure 7). . . 86 11. Plot of canonical axes 3 and 4 showing ordination of specimens according to population codes of collection sites (Table 5) (Compare with Figure 8). . . 87 12. Plot of canonical axes 1 and 2 ordinating specimens of Carex brevior, Carex molesta and Carex festucacea by whether they were collected at allopatric sites or sympatric sites (Table 5). Lines delimit boundaries of intermediate specimens (Compare with Figure 6) ...... 88 1

CHAPTER 1

INTRODUCTION

The same person who has made up his mind that the grasses are very difficult to identify is pretty apt to consider the sedges almost impossible. (Harrington 1977)

Carex is the largest genus within the Cyperaceae, with an estimated 2,000 species. They have a worldwide distribution, but are predominately found in temperate and boreal climates. Mackenzie (1931-1935) listed over 500 species for

North America alone. Eilers and Roosa (1991) list 108 species of Carex in Iowa, making it the largest genus in the state. Members of the genus Carex ( carices) are found thoughout Iowa in nearly every habitat including residential yards, but are more prevalent in wetlands meadows, river sloughs, marsh borders and fens.

The most troublesome Iowa sedges are Carex brevior, _c. fesmcacea, _c. molesta, .C. normalis, _c. tenera and _c. tenera var. echinodes (Gilly 1946). These form a species complex which I informally call the _c. brevior group (Table 1). These six taxa are part of an even larger aggregate of species associated with _c. straminea, whose morphological features intergrade into one another. Botanists have had difficulty classifying members of this aggregate for nearly two centuries.

The objectives of my study were: 1) to randomly sample up to 30 specimens from each population, at several populations of each taxa, to obtain statistical parameters for morphological characters and ascertain if variation occurs within populations, between populations, and between taxa; 2) to evaluate

44 characters and 12 character ratios for each taxa by Univariate Analysis and

Stepwise Discriminant Analysis (SDA) and obtain reliable taxonomic characters 2 or character combinations; and 3) to test the validity of taxonomic classifications within the ..C. brevior group by determining if the morphological forms are significantly different using Canonical Discriminant Analysis (CDA).

Table 1. The Iowa taxa of the Carex breyior group ( Cyperaceae ). Code Taxon B Carex brevi or (Dewey) Mackenzie

F Carex festucacea Schkuhr ex Willdenow

M Carex molesta Mackenzie ex Bright

N Carex normalis (Dewey) Mackenzie

T Carex tenera Dewey

E Carex tenera var. echinodes (Fernald) Wiegand

Statement of Problem

Carices are often difficult to classify because of their unique morphology and confusing taxonomic history. Recognition of taxa requires fruits which are mature but not overripe. The perigynia are frequently small, sometimes 2 mm or less in length, and features of the perigynium such as nerves and serrations are important. Accurate measurements of the perigynia are crucial (Voss 1972).

Individual taxonomic characters overlap among the taxa and cannot be relied upon with certainty. This, combined with a large number of similar taxa, makes accurate taxonomic keys arduous to construct and use. Many carices were 3 described before typification of taxa was common practice. In many cases, early authors of taxonomic names did not fully comprehend the boundaries of their classifications. They either included morphological forms of what later became separate species or described only one of the many forms of a particular taxon.

Some caricologists placed too much emphasis on a single character for identification. Since Willdenow (in Schkuhr 1801} first described _c. straminea there have been no less than 22 treatments involving this species and its allies, including the ..C. brevior group. During their careers, noted Carex monographers

L H. Bailey, M. L Fernald and K. K. Mackenzie interchanged names and taxa, e.g., nearly all the currently accepted taxa of the ..C. breyior group carried the name _c. straminea at one time, either as the perceived "type" form or as one of its varieties. Past and present difficulties associated with Carex taxonomy, especially the ..C. brevior group, can be attributed to nearly two centuries of confusion while botanists struggled to separate taxa through studies of morphology, genetics and ecology in an effort to formulate modem species concepts.

Tuckerman (1843) and many other 19th century botanists classified microspecies of the ..C. straminea aggregate as varieties of ..C. straminea. L H. Bailey (1883) called _c. straminea and its allies one of the six most troublesome groups of carices in North America. Bailey ( 1885) wrote, ·~. straminea is remarkable from the fact that all its varieties are connected with the type by a complete series of gradations. The individuals of these intermediate forms are also common." Boott (1862) wrote, "I believe that any one patiently studying the ----

4 group from ample material will be obliged to admit that it is impossible to discover exclusive characters on which any satisfactory specific distinctions can be found."

K. K. Mackenzie (1931-1935) described 533 species of Carex in North

America alone with very few varieties. He restored the taxa of the .C. straminea group to species status, often using highly variable morphological features of the inflorescence, leaf sheath and perigynia. Modem morphological studies of other groups have shown that the characters used by Mackenzie for species distinctions were among the most variable (Reznicek and Ball 1974; Waterway 1990).

Fernald (1942) was so exasperated with Mackenzie's choice of taxonomic characters that he wrote, "Suffering for many years from abnormal vision,

[Mackenzie] thought he saw ... what some others could not detect." Gilly (1946) wrote in his monograph:

...the section Ovales comprises one of the most difficult species­ groups of the genus Carex. I believe that Mackenzie, and most other modern workers as well, have recognized entirely too many species in this section of the genus. Because of variation among individuals of single colonies, the value of certain characters for the identification of and recognition of species may well be questioned.

M. L. Fernald, who began his long botanical career as a Carex monographer, recognized many of the species in question (Fernald 1950), but challenged the taxonomic characters used to separate them (Fernald 1942).

Specimens of the .C. breyior group can be categorized into characteristic morphological forms which reoccur throughout regions of North America. These forms have been described under various names in several historical publications.

The question becomes: are these forms separate species or polymorphic forms of 5 a single species?. If they are distinct species then reliable characters are necessary to deliminate and distinguish them. To ascertain the reliablity of taxonomic characters, it is necessary to delimit their ranges of variation. The classification of the ,C. breyior group, and other similar groups of sedges, binges on whether taxonomic characters vary because of genetic differences or enviromental influences.

Genetic studies of species groups have indicated that there are low levels of genetic variation within populations and that most of the genetic variation within species occurs between populations (Wbitkus 1988; Bruederle and

Fairbrothers 1986; Waterway 1990; Bruederle and Jensen 1991). Therefore, it is likely that variation within populations can be attributed to environmental influences (Smith 1967 1969; Smith and Faulkner 1976).

The morphological forms which Mackenzie and others describe are discemable if not entirely distinct for the .C. brevior group. These forms consistently reoccur in discrete habitats throughout their geographic distribution.

Herbarium studies alone can not determine if variation can be attributed to overlapping taxa or a single polymorphic species (Gilly 1946). Extensive morphological study of each taxon is required at the population level, at multiple sites, to determine if variation occurs within populations or between them (Sokal

and Rohlf 1969; Sneath and Sokal 1973). Univariate statistics of randomly

collected samples can determine character parameters and multivariate analysis

can evaluate taxonomic classifications using several characters in combination

even though individual characters overlap among taxa (Sneath and Sokal 1973; 6

SAS Institute Inc. 1988). Random sampling techniques are required to remove research bias in the selection of morphological forms for statistical analysis and allow statistical inferences to be made about populations of Iowa taxa (Sokal and

Rohlf 1969).

Taxonomic Classifications

The Genus Carex

Members of the genus Carex ( Cyperaceae) are usually not appreciated for their spectacular floral displays, because Carex is comprised of perennial grass­ like species with highly reduced, wind pollinated flowers. Nonetheless, a few species such as .C. ~ Carey have inflorescences large enough to be noticed and admired. Peattie (1939) commented that among his colleagues at the U. S.

Department of Economic Botany, sedges were most noted for their uselessness.

In reality, they are a significant forage for livestock in alpine meadows, steppes, and tundra regions of western North America, northern Europe, USSR and

Iceland (Catling et al. 1990). While some species are serious weeds, others have potential applications for controlling erosion and trapping nutrients.

Mm:pholoey. Members of the genus Carex ( carices) have unisexual flowers subtended by a scale-like bract. The flowers are adapted for wind-pollination and lack vestiges of the perianth found in other sedges, such as bristles or scales.

Carices are recognized by a unique structure called the perigynium, a sack-like leaf which completely encloses the pistillate flower and resulting achene. The ovary of the pistillate flower is either lenticular or trigonous with 2 or 3 styles.

The perigynium and subtending scale is commonly refered to as the female floret. 7

However it is more accurately termed a spikelet, because inside the perigynium, and ventral to the pistillate flower, is an aborted axis refered to as the rachilla.

Anatomical evidence suggests that the pistillate flower, perigynia and rachilla are derived from an ancestral, multiflora! spike (Smith 1966; Smith and Faulkner

1976). The male floret has 3 stamens subtended by a scale-like bract. The vascular arrangement of the staminate floret suggests that it was derived from three, uni-staminate flowers (Smith 1966). Staminate and pistillate structures and their subtending scales will be termed "florets" in the vernacular sense for purposes of this discussion. The florets are arranged in spikes which are either unisexual or bisexual.

Subceneric classifications. The various arrangements of florets and spikes in the inflorescence provides much of the basis for sub-generic classification within Carex. Gilly (1950 1952) described 111 inflorescence types in 988 species of ~- There are 3 subgenera of Carex from which 71 sections are described for North America (Mackenzie 1931-1935). Within the sections are subsections, which are further divided into aggregates of morphologically similar species.

Species have often been sub-classified into varieties and forms.

Kukenthal (1909) described four subgenera within Carex, of which,

Rezincek (1990) recognizes Carex (Eucarex), Indocarex and Vicnea. A fourth subgenus, Primocarex Kukenthal, is not recognized because it is thought to be an artificial conglomeration of taxa with only a single spike in the inflorescence.

The genus Carex has been further subdivided into sections and subsections.

Tuckerman (1843) was the first to devise a natural classifications system for Carex 8 apart from the Linnaean system of subgenera (Gray 1843; Robertson 1979).

Tuckerman (1843) proposed five sections which he further divided into subsections. Species were grouped into analytical categories and aggregations of morphologically similar species which were treated as varieties of a base species, such as ,C. straminea. Bailey's (1886) classification system included subgenera,

Tuckerman's sections, and subsections (which included the section Ovales).

Kukenthal (1909) wrote the most recent worldwide treatment of the genus Carex.

His system of subgenera and sections are still in use today (Mackenzie 1931-

1935; Fernald 1950; Gleason and Cronquist 1963; Voss 1972, etc.). Mackenzie

(1931-1935) created sub-sectional divisions, but never specified their rank. He gave each subdivision a name based on a representative species (e.g., ,C. festucacea for his subdivision Festucaceae). Hermann (1974) ranked Mackenzie's subdivisions as subsections. I have accepted Mackenzie's Festucaceae and

Tribuloideae as subsections, mainly for convenience, but I do not know if these are accepted as nomenclaturally valid. Whitkus (1981) claims they are not.

Recent studies have attempted to define taxa within aggregates or complexes of morphologically similar species (Reznicek and Ball 1974; Whitkus and Packer 1984; Standley 1985 1987; Rettig 1986; Crins and Ball 1989;

Bruederle et al. 1989). I have used the term, "species aggregate" as defined by

Davis and Heywood (1973) to describe ,C. straminea and its allies. These form a group of component species, which are taxonomically distinct and presumably

closely related but difficult to distinguish. Microspecies within species aggregates tend to have fewer distinguishing characters than other species of the genus. But 9 it remains to be determined if taxonomic characters of the Carex brevior group are constant and whether the species are effectively isolated from one another.

The Subienera of Carex

Subgenus Carex generally has unisexual terminal spikes. The lateral spikes are either unisexual or androgynous (monoecious spikes with superior staminate florets). The spikes are mostly peduncled, but occasionally sessile. The ovaries are mostly tri-stigmatic, but sometimes bistigmatic. These result in trigonous or lenticular achenes, respectfully. The perigynia are usually terete or trigonous but a variety of shapes are known. Carex is the largest subgenus with 1,400 species distributed throughout the world (Reznicek 1990).

Subgenus lndocarex has bisexual, androgynous spikes (superior staminate florets) and tristigmatic achenes. The perigynia are trigonous or somewhat terete.

There are about 100 species found primarily in the tropics and subtropics of southeast asia, but also in the Paleotropics and Neotropics (Rezinecek 1990).

Subgenus Vi~nea. which includes the .C. brevior group, has bisexual spikes which are usually sessile on the rachis of the inflorescence. The lenticular achenes are usually bistigmatic. Most spikes are androgynous but at least 5 sections, including the Ovales, are gynaecandrous with superior female florets on

the bisexual spike. Vignea carices are found mainly in North and South America

and in the temperate and boreal regions of Eurasia with some representatives in

the Paleotropics (Rezinicek 1990). 10

The Section Ovales Kunth.

The section Ovales has the largest number of species and is considered the most difficult among North America carices (Mackenzie 1931). Gilly (1946) listed 15 species of the section Ovales in Iowa (Table 2). These are recognized by gynaecandrous spikes with female florets located above the male florets. The perigynia are flat and winged at their margins. The achene is plano-convex or

lenticular (biconvex). The following is taken from Mackenzie (1931-1935):

[Rhizomes] densely caespitose, or with more or less prolonged rootstocks; culms triangular, hollow; leaf-sheaths not red-dotted nor cross-rugulose ventrally, [hyaline] but sometimes green-striated; spikes from 2 or 3 up to 20, with several to many perigynia, the terminal gynaecandrous, the lateral pistillate or gynaecandrous, simple, the inflorescence varying from capitate to moniliform; lower bracts from inconspicuous to very conspicuous; perigynia varying from scale-like or flat (except where distended by achene) to thick and strongly plano-convex, the body subulate to reniform, narrowly to broadly wing-margined, appressed or ascending or spreading, little corky-thickened at base, promimently beaked, the beak sutured dorsally, bidentate, or obliquely cut, usually becoming bidentulate or bidentate, usually serrulate on the margins, rarely smooth; achenes lenticular, apiculate, jointed with the straight, slender style; stigmas 2.

Table 2. Taxa of the Care~ section Ovales in Iowa (Gilly 1946). Carex bebbii Olney, Carex bicknellii Britton Carex brevior (Dewey) Mackenzie Carex cristatella (Dewey) Britton Carex festucacea Schkuhr ex Willdenow ~ molesta Mackenzie ex Bright Carex muskingumensis Schweinitz Carex normalis (Dewey) Mackenzie Carex scoparia Schkuhr Carex projecta Mackenzie Carex suberecta (Olney) Britton ~ sychnocephala Carey ~ tenera Dewey ~ tribyloides Wahlenberg 11

The Subsection Festucaceae Mackenzie (nomina nuda).

Mackenzie's subdivisions were treated as subsections by Hermann (1974), but it is not certain if they were ever validly published. Mackenzie (1931-1935) provided a description to the Festucaceae Mackenzie in a dichotimous key to his

Ovales subdivisions which are repeated here in text form:

Bracts not leaf-like nor conspicuously exceeding [the inflorescence]; perigynia with beak flattened and margined at tip, serrulate to the apex, often bidentate as well as obliquely cut dorsally; scales shorter than the perigynia and noticebly narrower above, largely exposing perigynia above; sterile culms [often poorly developed, leaf-blades erect or ascending, usually clustered at apex]; perigynium-body not obovate, widest near the middle or base.

The Carex breyior Group

The following synopsis of common characteristics of the .C. brevior group was adapted from Mackenzie's (1931) descriptions of .C. brevior, .C. festucacea, C. molesta. _c. normalis, and _c. tenera:

Ve2etatiye characters: roots densely cespetose; rhizomes short, black and fibrillose; fertile culms 3-10 (15) dm long, slender to base, stiffly erect, ascending or nodding, usually exceeding the leaves, sharply triangular and roughened on the angles beneigth inflorescence, culm base brownish to black and clothed with scenescent leaves, lower nodes not exposed; leaves well developed, either 3-6 or

4-7 regularly distributed on the lower 1/3 to 1/2 of the fertile culm; leaf blades light-green or yellow-green, flat, V-shaped or flanged V-shaped (inversely W­ shaped) (terms follow Metcalfe 1971), thin or thick, erect, ascending or spreading,

0. 75 to 4 dm long, 1.5-2.5 mm, either 2-4 mm or 4-6 mm wide, rough to very rough on margins; sheaths tight (loose on .C. normalis), dorsal side sometimes 12 septate-nodulose with green or green and white mottled coloratio~ ventral surface conspicuously white-hyaline, prolonged beyond base of blade and continuous with ligule; vegetative cu1ms conspicuous, with ascending or spreading leaves bunched at apex.

Inflorescence characters: spikes (inflorescence units), gynaecandrous, 3-10, varying from aggregate, aggregate to spreading at base, moniliform (spike tips approximate to spike bases}, or strongly moniliform arrangements, rachis flexuous to stiffly erect, 2.5-10 in length, 7-15 mm in width; varying from subglobose to obovoid, 6-16 mm long, 4-8 mm wide, apex varing from blunt to tapering, spike­ base varying from truncate, rounded, tapering to long-clavate (lateral spike bases sometimes differing from terminal bases); staminate florets few to numerous, ranging from inconspicuous to nearly half the spike; inflorescence bracts either short and scale-like or with a setaceous bristle of varying length; scales ovate, apex obtuse, acute or short accuminate, green-hyaline with 3-nerved center

(golden yellow at maturity}, scale width narrower than or equal to perigynium width, scale length shorter than or exceeds perigynium beak.

Periwum characters: perigynia 10-30 per spike, appressed, ascending or widely spreading, plano-convex or slightly concavo-convex, subcircular, broadly ovate or narrowly ovate, rounded at base, tapering, constricted to abruptly narrowed to beak tip, 2.5-6 mm long, 1-4 mm wide, narrowly or broadly winged to base, serrulate above middle, membraneous or coriaceous, faintly to strongly 5-7 nerved on dorsal surface over achene, faintly to strongly (0-7) nerved on ventral surface over achene (sometimes strongly nerved at perigynium base}, some 13 species with nerves in the wings, usually green-white becoming brown or straw­ colored at maturity; beak length 1/4 to 1/2 perigynium length, beaks are obliquely cut at apex, bidentate, with dorsal (abaxial) suture.

Achene characters: achenes ovate, elliptical to oblong, 1.25-2 mm long, 1-

1.75 mm wide, apiculate, sub-stipitate to stipitate at base, yellow to brown; styles straight (sometimes bent at base), jointed with achene, deciduous, 2 stigmas.

The Taxonomic Histocy of the Carex brevior Group.

The ,C. brevior group is an informal name for 5 species and one variety which resemble one another and presumably are closely related. The morphological forms intergrade into one another and are part of an even larger

aggregate of North American carices which Boott (1862) called·~. straminea and its allies." Specimens of the ,C. brevior group have been closely associated with

the name ,C. straminea. There have been at least 22 different taxonomic

treatments which included the Carex straminea aggregate. These treatments

either classified taxa within the aggregate as varieties of .C. straminea or as distinct species. L H. Bailey (1885) combined seven currently recognized taxa

under the name ,C. straminea without describing any varieties, while Olney (1870)

created numerous varieties and forms in an effort to descriptively document

variation within the aggregate. In addition, there has been ambiguity about which

form or taxon was represented by the type of Carex straminea. There were

periods of time when specimens of either Carex brevior or Carex tenera were

thought to be the "typical" form of Carex straminea. Treatments of these periods

describe and illustrate Carex brevior or Carex tenera as Carex straminea. What 14 follows is a brief description of the taxonomic history for each taxa of the Carex breyior group and Carex straminea. A referenced nomenclator of the .C. brevior group and .C. straminea is provided in Appendix A

Taxonomy of Carex straminea

The taxonomic problems associated with .C. straminea began with the

original description by Willdenow in Schkuhr's (1801) monograph. According to

A A Reznicek (pers. comm.), Willdenow received the Reverend Klaproth's

specimen from North America probably before 1800. Willdenow named it .C.

straminea and sent it with a short description to Schkuhr. Willdenow's original

description of the perigynium was "capsulis obovatis acuminatus" and Schkuhr's

illustration (1801: Tab. G No. 34) shows an obovate perigynium with a short

acuminate beak. No specimen filed under .C. straminea in Willdenow's

Herbarium in Berlin nor in Schkuhr's Herbarium in Halle fits this description or

illustration. Willdenow (1805) changed his description of .C. straminea in the

fourth revision of Linneaus' Species Plantarum. The perigynium was then

described as "fructibus subrotundo-ovatis rostratis bidentatis", i.e., the perigynium

subcircular to ovate, provided with a long, bidentate beak. Schkuhr (1806)

repeated Willdenow's second description in his revision and illustrated (Fig. 174)

the perigynium of Klaproth's specimen No.17177 to show a round body and a

long narrow beak. Schkuhr annotated the herbarium sheet of .C. straminea at

Halle with "the neck of the capsule in No.34 is drawn somewhat too short"

(Reznicek pers. comm. ). The confusion over the true identity of .C. straminea

lasted until Svensen (1938) fixed the type to Klaproth's specimen in Willdenow's 15

Herbarium. In the interim, several specimens of other taxa were erroneously identified as _c. stramine'h and the name was attached by various botanists to specimens of _c. breyior (Dewey) Mackenzie, _c. tenera Dewey and _c. albolutescens Schweinitz (Boott 1862; Bailey 1889; Mackenzie 1922).

Besides the type specimen of _c. straminea collected by Klaproth, there are specimens of other taxa attached to the herbarium sheets of Schkuhr and

Willdenow. Most are too immature and fragmented for positive identification by

Dr. A A Reznicek (pers. comm). Schkuhr's specimens were received from

Reverend Muhlenberg: one is ..C. cristatella (Dewey) Britton; the other resembles

.C. bebbii (Bailey) Fernald or _c. normalis (Dewey) Mackenzie. Attached with

Willdenow's type of _c. straminea is another specimen, possibly _c. tribuloides

Wahlenberg. Also, Schkuhr's herbarium sheet of _c. straminea has a fragment packet containing perigynia of Willdenow's type (Reznicek pers. comm.). I have examined Muhlenberg's herbarium at PH and found specimens of _c. cristatella and _c. normalis filed under the name of _c. straminea.

Mackenzie (1915) finally sorted out the true identity of _c. breyior, _c. tenera and _c. festucacea, but he misinterpreted specimens of _c. albolutescens as

.C. straminea. Mackenzie (1922 1931 1940) viewed the two Schkuhr illustrations of ..C. straminea as variants of ..C. albolutescens. Initially, Mackenzie (1915) wrote that Schkuhr's (1801) illustration was the true form of .C. straminea and that

Schkuhr's (1806) illustration depicts _c. straminea var. brevior Dewey. Later,

Mackenzie (1922) declared _c. albolutescens a synonym for _c. straminea sensu

Mackenzie (1915). Mackenzie's (1940) figure 184 of _c. straminea depicts a 16 perigynium taken from a specimen of .C. albolutescens. The body of this perigynium is nearly oval and closely resembles Schkuhr's (1806) figure 174.

Svenson (1938) discovered that the name .C. straminea should be applied to specimens known by .C . .ridtii Mackenzie. This in turn required the name .C. albolutescens Schweinitz be revived for the taxon treated as .C. straminea by

Mackenzie (Rothrock 1991).

The Iaxonomy of Carex brevior

Many specimens of .C. brevior were annotated, described, and/or illustrated as the "typical form" of .C. straminea by Torrey (1836), Carey (1856),

Boott (1862), and Bailey (1886). I attribute the erroneous association of ,C. brevior and .C. albolutescens with the name .C. straminea to Chester Dewey, who serially published his monograph on the genus Carex from 1818 until his death in

1867. In Dewey's (1826a) original description of ,C. straminea var. breyior, he wrote, "It was this variety which was described by Willdenow and to which the

name was given." Dewey also suggested that .C. albolutescens is a variety of ,C.

straminea. Late in his life, Dewey (1867) annotated Schweinitz's isotype (GH) of

.C. albolutescens as a synonym for his variety brevior. .C. albolutescens was initially described by Schweinitz ( 1824) in a leg of a

key, but it never appeared in Schweinitz's (1826) monograph edited by John

Torrey. Bailey (1893) restored the name .C. albolutescens to specimens annotated

by Torrey as ,C. straminea var. foenea ( = .C. longii) (In Muhlengerg's herbarium

(PH), I found specimens labeled Carex foenea Muhlenberg to be a synonym for

.C . .lmWi Mackenzie). 17

Dewey (1826a) cites Wahlenberg (1803) as the source of the primary description for .C. straminea Willdenow. Wahlenberg (1803) described the perigynium of .C. straminea as being subcircular to obovate. Many forms of the perigynia of .C. breyior are subcircular (Boott 1862). Typically, the perigynia of

.C. albolutescens is obovate (Rothrock 1991), but it also has orbicular forms

(Mackenzie 1931).

The Taxonomy of Carex tenera

Bailey (1889; 1890) mistakenly associated Willdenow's type of .C. straminea with specimens known by Carey, Sartwell and Olney as .C. tenera Dewey ( = .C. hounathodes). Following this, specimens of the typical form of .C. tenera Dewey were named .C. straminea by Mackenzie (1896 1913), Fernald (1902 1908) and

Kukenthal (1909). Specimens of .C. hormathodes Fernald were mistaken for C. tenera Dewey by Sartwell, Olney, Bailey (1883; 1885), Mackenzie (1896; 1913) and Fernald (1902). In fact, Chester Dewey facetiously annotated specimens of

,C. hormathodes as .C. tenera Olney not Dewey, essentially accusing S. T. Olney for the origin of the error. Much chagrined by this, Olney wrote a long annotation claiming he was "surprised on receiving Boott's (1862) illustration to find myself quoted under these plants as .C. tenera Olney" ( GH). Kukenthal

(1909) cited illustrations of .C. hormathodes as depicting .C. tenera Dewey.

Fernald (1906) distinguished Dewey's "original" specimens of .C. tenera Dewey

(GH) from specimens of .C. hormathodes (Boott) Fernald; meanwhile, Dewey's type specimens of .C. tenera were classified as .C. straminea until Mackenzie

(1915). Mackenzie (1931-1935) designated Dewey's original specimens of .C. 18 tenera as lectotypes. The best illustration of Dewey's original ,C. tenera is Boott's

{1862) figure 384.

The Taxonomy of Carex tenera yar. echinodes

Fernald (1902 1950) described ,C. tenera var. echinodes Fernald as having

"tips of the slightly longer perigynia divergent and conspicuous." Kukenthal

{1909) reduced its classification to forma echinodes. Neither classification is recognized by Mackenzie or later authors. Mackenzie's (1940) illustration of the perigynium of ,C. tenera appears similar to variety echinodes with a relatively narrower perigynium and slightly longer beak than Dewey's type. The illustrations of Gleason (1952) and Voss (1972) were taken from Mackenzie

{1940). Many other authors have used these illustrations.

The Taxonomy of Carex festucacea

The taxonomy of ,C. festucacea Schkubr ex Willdenow is equally as confusing. The original description of ,C. festucacea by Schkubr appeared in

Willdenow {1805). Later, Schkuhr (1806) illustrated _c. festucacea in figure 173. The type specimens of ..C. festucacea are missing, however, Schkubr's illustration fixes the application of the name (Rothrock 1991). Dewey (1824 1836) placed too much emphasis on clavate spike bases for the recognition of ..C. festucacea.

Despite Torrey's (1836) warning that this character was unreliable, many later botanists used the clubbed-shaped spike base as the only criteria for recognizing

,C. festucacea. Consequently, specimens of several taxa were identified as ,C. festucacea (see Boott 1862). 19

Somehow the name .C. festucacea became associated with specimens currently named .C. merritt-fernaldii (Bailey 1889, Mackenzie 1896 1913 and Fernald 1902 1908). Fernald (1902) wrote: "Schkuhr's .C. straminea of figure 174 which we now know to be different from Willdenow's plant of that name, was an extreme form of .C. festucacea (=.C. merritt-fernaldii)." Both Fernald (1902 1908) and Mackenzie (1896 1913), described and illustrated specimens of .C. merritt-fernaldii Mackenzie under the name .C. festucacea (Mackenzie 1922).

Fernald (1902 1908) classified specimens of .C. brevior as .C. festucacea var. brevior (Dewey) Fernald.

The Taxonomy of Carex normalis

Specimens now known as .C. normalis (Dewey) Mackenzie have been well defined and easily recognized since Dewey's (1836) original description as .C. mirabilis. However, there have been several different varietal synonyms assigned to the taxon reflecting the various interpretations of its relationship to other species: Tuckerman (1843) classified it as a variety of .C. straminea; Boott (1862) saw it as a variety of .C. cristatella; and Olney (1870) treated it as a variety of .C. tribuloides. The morphology of .C. normalis is intermediate between .C. tenera and .C. tribuloides. These species represent two distinctive morphological aggregates of species recognized by Mackenzie (1931-1935) as subsections

Festucaceae and Tribuloideae.

Fernald (1902) described ".C. normalis" var. perlon2a which he later reduced to a forma perlon2a (Fernald 1950). 20

The Taxonomy of Carex molesta

Mackenzie (1931-1935) described .C. molesta and distributed isotypes from

Quindaro, Wyandotte County, Kansas. However, Bright (1930) previously published the name .C. molesta Mackenzie with a description of a specimen from

Pennsylvania. Rothrock (1978) has cited this specimen as the type and declared the proper name to be .C. molesta Mackenzie ex Bright. Bright's isotype (PH) is the same species as Mackenzie's isotypes (KSU). Gates (1940) classified the taxon as .C. brevior var. molesta. Gleason (1952) and later Cronquist (Gleason and Cronquist 1963) considered the taxon as part of .C. brevior or a putative hybrid created by .C. brevior x .C. noonalis. Both Fernald (1950) and Voss (1972) recognized .C. molesta as a distinct species.

Carex Evolution. MorphololO' and Genetics

Pbylo~enetic Development of the Inflorescence

Evolutionary relationships among and within the genera of the tribe

Cariceae (Care~ Kobresia, Unicinia, and Schoenoxiphium) are poorly understood

and probably remain so until generic and subgeneric relationships are well

defined (Reznicek 1990; Crins 1990). Evolutionary hypotheses for the Cariceae

are based primarily on developmental patterns of the perigynium and

inflorescence among the genera (Gilly 1950 1952; Nelmes 1952; Smith and

Faulkner 1976; Rezincek 1990). It is generally assumed that Carex species with

monospicate inflorescences are derived from precursors with multiple spikes on

highly branched inflorescences through a series of reductions in a manner similar

to Zimmerman's (1930) theory of organogenesis. The opposing hypotheses best 21 espoused by Smith and Faulkner (1976) and Rezinicek (1990) differ on which subgenera of Carex have the primitive or the advanced form of the inflorescence.

While the interpretations differ, both hypotheses agree that the diverse forms of the inflorescence, caused by developmental differences in inflorescence meristems, reflect the evolution (or phylogeny) of the tribe. Morphological development explains the great diversity of inflorescence types among the subgenera, but it also provides insight to the origin of morphological variation expressed by species.

The phylogenetic interpretation of the perigynium is widely disputed, as witnessed by the 18 names recorded for it by Holm (1896). In contrast to its interpretation as a prophyll, it has been considered as a reduced bract, a pericarp, a nectary or disk, a perianth and a utricle. At one time the perigynium was held to be homologous with the palea of the Poaceae and defined as a single bract with its margins fused into a suture on the abaxial or dorsal side (Townsend

1885). Snell ( 1936) considered it to be a prophyll, or the first leaf of a lateral axis in monocots. In Carex. the prophyll is reduced to a bladeless sheath. Smith and Faulkner (1976) hypothesized that the perigynium was derived from another type of prophyll once called an ochrea (Townsend 1885) but now called a cladoprophyll (Holm 1896). The cladoprophyll is a bladeless leaf sheath surounding bases of inflorescence branches (i.e., peduncles of lateral spikes) of subgenera lndocarex and Carex (Holm 1896; Snell 1936; Holttum 1948; Blaser

1944; Smith and Faulkner 1976). Cladoprophylls are often hidden inside the sheath of a subtending leaf or bract and are either sterile or fertile. Holm (1896) 22 distinguished the fertile cladoprophylls as "anthroprophylls". There is a third type of prophyll sessile to spike bases of carices within the subgenus lndocarex.

Reznicek (1990) describes these as "perigynium-like prophylls" and used the term

"inflorescence prophyll" to distinquish them from cladoprophylls.

Anatomical studies of the vascular bundles of the cladoprophyll and the perigynium support the conclusion that they are reduced leaves. Both their vascular morphology and relative position are that of a leaf sheath of Carex

(Snell 1936). In addition, morphological studies have shown there is an aborted rachilla or spikelet axis at the ovary base within the perigynium (Snell 1936;

Smith 1966). In some species of the tribe Cariceae the rachilla is prolonged and exerted beyond the apical orifice of the perigynium. Therefore, the combined structures of the pistillate flower, rachilla, perigynium and subtending scale, are not actually a single "floret" but a reduced multiforal spikelet in the base of a reduced bract (Snell 1936; Svensen 1972; Rezinecek 1990). The ovary is axillary to an aborted spikelet axis and the scale is analogous to the lower glume of grasses. In addition, the vascular system of carices indicates that the so called

"male florets" of the inflorescence are actually derived from at least three male flowers of a former spikelet (Smith 1966).

Smith and Faulkner (1976) believe that Carex prophylls were derived entirely by a process of reduction: 1) the cladoprophyll was created when the lamina was lost, leaving a tubular sheath enclosing the bases of spike peduncles;

2) the development of the rachilla ceased leaving a spike with a solitary ovary

(floret); 3) the peduncle supporting the derived "floret" diminished, leaving the ovary and the aborted rachilla inside a tubular "perigynium". 23

In contrast, Reznicek (1990) hypothesizes independent evolution of the cladoprophyll and perigynium. Reznicek believes the two prophylls were derived by a combination of reduction and proliferation of the rachilla: 1) an ancestral species developed a perigynium precursor which later diversified into what became the tribe Cariceae; 2) cladoprophylls in the subgenus Carex originated from empty pistillate scales at the bases of spikes; 3) the inflorescence prophylls

of Indocarex were derived from a perigynium whose rachilla proliferated into the

characteristic multiflora! spike of lndocarex. Reznicek states his theory is more

inclusive of tropical species of Carex, while other theories were based mainly on

temperate species.

It is not within the scope of this paper to fully discuss the different

hypotheses: Reznicek (1990) does that quite adequately. However, a discussion

of the morphological development of the inflorescence is necessary to understand

taxonomic relationships within species aggregates. On the basis of morphological

development of the apical meristem in Carex and related genera (tribe Cariceae ),

it is generally accepted that: 1) any inflorescence of Carex can be interpreted as a

repeatedly branching system in which each ultimate branch either develops into a

flower or aborts; 2) apart from abortion, Cariceae meristems have three possible

outcomes: they may become i) male flowers, ii) female flowers, or iii) develop

into compound structures, namely spikes; 3) the growth of a male flower

primordium is determinate, but growth of a spike primordium is indeterminate

and it may develop into anything from a one-flowered female spikelet to a

branched bisexual spike; 4) the differences in inflorescence morphology which 24 characterize the subgenera of Carex are explicable in terms of the relative degree in development of the meristem at female flower nodes (Smith 1966; Smith and

Faulkner 1976).

Pbenot)llic Plasticity and Meristematic Development The morphology of the inflorescence varies considerably within the genus

,Cmn. Gilly (1950 1952) described 111 inflorescence types in the 988 species he examined. As described above, each of these inflorescence types are derived from three possible outcomes of the meristem, indicating that genetic mutations alter the morphological arrangement of the inflorescence. Large morphological differences between suprasectional taxa may have resulted from an accumulation of gene mutations responsible for inflorescence development. Minute differences of the inflorescences observed within species aggregates may be attributed to a smaller number of developmental alterations. However, morphological development due to genetic differences may be moderated by the environment.

Meristems of Carex are affected by hormonal imbalances created by environmental stress which affect the regulation of genes (Smith 1967; Smith

1969; Smith and Faulkner 1976).

Phenotypic variation within species may be attributed to the substitution of any of the possible meristematic outcomes which can occur in the development of the inflorescence. Inflorescence development is controlled by plant hormones.

Smith (1967) applied various auxins and cytokinins during and after the transition to the flowering stage. He found that auxins: 1) reduced the number of inflorescences produced while maintaining the number of florets per 25 inflorescence; 2) increased the number of lateral spikes; 3) induced the production of female florets in potential male sites; and 4) increased the ratio of female florets to male florets in the inflorescence. Kinetins applied continuously throughout the growing season completely suppressed the development of the inflorescence. Specimens treated with 3 weekly treatments exhibited: 1) increased branching in vegetative shoots; 2) reduced numbers of inflorescences and reduced

numbers of florets per inflorescence; 3) reduced the height and internode length

of the inflorescence, and 4) suppressed branching of the inflorescence. Smith

(1969) observed similar morphological responses in specimens where leaves and

roots were removed. Removal of leaves upsets initiation and growth of the

inflorescence and Smith {1969) concluded that the continued stimuli from leaves

is essential for normal development. Removal of roots or root apices disrupts

inflorescence initiation and branching, and Smith (1969) concluded that actively

growing roots is essential for normal branching to occur. He suggested that

normal branching of the inflorescence may depend on an adequate supply of

cytokinin from the roots. Smith and Faulkner (1976) discuss an example of

inflorescence abnormalities produced in .C. flacca which had been trampled by

cattle (while dormant) during a wet winter. The abnormalities discribed were

similar to those observed by the application of hormones (Smith 1967). Smith

and Faulkner (1976) suggested that sex expression within Carex inflorescence can

be explained by physiological gradients controled by environmental conditions. 26

Genetics

Cyto2enetics. The unique cytogenetic structure of Carex accounts for both the large morphological diversity between subgenera as well as the morphological similarity of microspecies within species aggregates. The centromere is diffused and not localized on any one portion of the chromosome (Davies 1956). Carex chromosomes have a sticky matrix causing the ends to agglutinate together to form a conglomerate chromosome network with an irregular outline.

Chromosomes readily fragment with a portion of the centromere. These pieces can be included in future meiotic divisions, either as separate entities

(chromosomes) or by reattachment (Davies 1956).

Wahl (1940) observed a reversal of meiotic divisions. The centromere splits, separating the chromatids during the first meiotic division; the second meiotic division is reductional. Wahl (1940) wrote, ''The same number of chromosomes were always found at both metaphase I and II while different and quite irregular numbers were frequently found in the microspore nucleus of a hybrid plant, or an individual with multivalents." Davies (1956) concluded that the reversal of these two divisions preserves chromosomal aberrations.

Agmatoploidy. Aneuploidy is common throughout Carex. Aneuploid chromosome series have been documented in subgenera, sections, species­ complexes, and within species (Hielbom 1928 1939; Tanaka 1940; Wahl 1940;

Davies 1956; Faulkner 1972; Whitkus 1981 1988; Whitkus and Packer 1984;

Hoshino 1981; Nishikawa et al. 1984; Standley 1985; Crins and Ball 1988). In the

~ species examined, there exists a series of chromosome numbers with 27 haploid numbers ranging from n = 6 ton = 56 (Heilbom 1939; Wah11940;

Davies 1956; Hoshino 1981). It was speculated that Carex exhibited a high degree of polyploidy, octoploid or higher (Wahl 1940). Heilbom (1939) considered the basic chromosome number was 7, because n = 28, 42, 56 occur in the literature most frequently. Chromosome counts not divisable by 7 were probably produced by aneuploidy. Mter tabulating chromosome counts of 305 species and 16 varieties, Tanaka (1949) proposed that base ploidy numbers 6, 8,

9, 10 and 12 were secondarily balanced from initial ploidy numbers 3, 4, and 5.

These researchers hypothesized that Carex evolution occurred initially by multiple increases in chromosome sets (polyploidy) followed by singular additions and deletions of chromosomes (aneuploidy) (Hielbom 1939; Wahl 1940; Tanaka 1940

1949).

However, Tanaka (1949) reported that no polyploid species had been found in the genus. Davies (1956) stated that autopolyploidy is exceedingly rare in the genus and plays a very small part in the evolution of Carex and she concluded:

The series [in Carex chromosomes] have almost certainly arisen in the first instance by chromosome breakage, agmatoploidy, and hence the species evolved through small and gradual changes in the course of time. It would therefore seem likely that the lower­ numbered species have given arise to the higher numbered, and consequently the series is ascending. However, the fact that the aneuploid series of numbers have arisen by fragmentation, explains why the species with the higher numbers of chromosomes have increasingly smaller chromosomes.

Davies (1956) hypothesis is supported by a more recent study of 50 taxa

(46 species and 4 varieties) in 21 sections of Carex (Hoshino 1981). Crins and 28

Ball concluded that agmatoploidy has been the dominate process of chromosomal evolution in the Carex. section Ceratocystis. Grant (1981) wrote that "fusion and fission of chromosomes (agamatoploidy) in Carex is the accepted hypothesis for chromosome number evolution in the genus, although strict aneuploidy cannot be ruled out."

Davies ( 1956) found chromosome morphology is a useful taxonomic character which is correlated with morphological arrangements of species within sections. Hoshino (1981) found chromosome races in different habitats and regions of geographic distribution of a species. Speciation is probably initiated by chromosome aberrations forming chromosome races which became geographically isolated. Whitkus (1981) hypothesized that the section Ovales diversified during the pleistocene as glacial ice retreated. He speculated that precursor species colonized the exposed glacial till. Through geologic time, the homogenous glacial landscape differentiated into discrete habitats or microhabitats, ecologically isolating chromosome races which later evolved into species.

Hybridization. Hybridization also played a role in the evolution of Carex

(Tanaka 1949; Whitkus 1988). Cytogenetic research in Carex involves using natural and artificial hybrids among closely related species. Heilbom (1928) counted chromosomes at meiotic stages of natural hybrids--specifically heterotypic metaphases in pollen mother-cells. Following meiosis, Heilborn observed variation in the number and size of chromosomes. Meiotic divisions of hybrids results in the formation of functional gametes with additional chromosomes. New chromosome numbers arise in hybrids through the formation of univalents in 29 meiosis. Tanaka (1940) made crosses in 13 species to yield 19 hybrid combinations. The artificial crosses were successful 37.3% of the time, and

Tanaka suggested that natural hybidization occurred frequently and contributed to the origin of aneuploidy.

Intraspecific hybrids are a common occurrance in the genus and Tanaka

(1949) reported that 90 out of 149 species examined had several karyotype numbers. Some species were found with as much as 6 different chromosome numbers in an aneuploid series. Hybrid karyotypes are found in the _c. brevior group. Wahl (1940) reported that ..C. festucacea has a trivalent karyotype (2n = 71) which is frequently heteromorphic (Tanaka 1949). Also, ..C. tenera has races with different chromosome numbers. The ..C. brevior group forms an aneuploid series (Wahl 1940).

Tanaka (1949) reported that intraspecific hybrids [n = 9 x n = 10 and n =

17 x n = 18] proved fertile with karyotypes composed of herteromorphic bivalent or trivalent chromosomes. Frequently, karyotypes were found with univalents derived from the duplication of chromosomes. Hybrid crosses with larger chromosome numbers [n = 19 x n = 22] varied from cell to cell resulting in differing counts of quadrivalent, trivalent, bivalent and univalent chromosomes

(Tanaka 1949). Many of the bivalent chromosomes where non-homologous or heteromorphic pairings. Such structural hybrids account for the production of the

aneuploidy in the genus Carex. They result from the duplication or the loss of a

few chromosomes through meiotic irregularity. Chromosome hybrids have

minimal reductions in fertility (Tanaka 1949; Davies 1955; Faulkner 1973). 30

Artificial hybridization studies in Carex have concluded that closely related species are easily crossed. Reproductive isolation is maintained in nature through differences in geographical distribution and habitat (Davies 1955; Faulkner 1973;

Hoshino 1981; Standley 1985; Whitkus 1988).

Tanaka (1949) documented morphological differences in seed shape which he attributed to extra chromosomes. Davies (1956) noted that chromosome races with differing karyotype numbers can be larger and more robust than most forms

of the species. Wahl noted that ,C. tenera (n = 27) was the larger more robust

form of the species.

Population i:enetics of species aggre~ates. Carex populations are

genetically uniform exibiting little if any heterozygosity and reproductive isolation

occurs mainly by selfing or in-breeding (Whitkus 1988; Bruederle & Jensen 1986).

Whitkus (1988) conducted artificial selfing and hybridization among species

and races within the ,C. macloviana D'Urv aggregate (C. macloyiana D'Urv. [n =

43]; ,C. preslii Steudel [n = 40,41]; and ,C. pachystachya Cham. ex. Steudel [n =

37,38,39,41]. The races exhibited no morphological differences except for the .C.

pachystachya n = 41 race which represents one extreme of the range of variation

within the species (Whitkus and Packer 1984 ). Isozyme studies showed that

members of this group exhibited little genetic diversity with very low levels of

heterozygosity and species and races are distinct in nature as no populations are

known where two races or species interbreed (Whitkus 1988). Whitkus found

that: 1) these plants are self compatible; 2) interracial and interspecific crosses

were significant; 3) crosses within races or species were more successful than 31 interracial or interspecific crosses; 4) there was equal to greater degree of success in selfing than in outcrossing; and 5) apomixis was not a significant occurrance.

Whitkus (1988) concluded that an autogamous mating system, one which favors selfing over out-crossing, would maintain reproductive isolation in mixed populations.

Similar conclusions were reached in isozyme studies of the .C. crinita Lam. complex (Bruederle and Fairbrothers 1986). The majority of genetic variation was distributed among taxa indicating species are highly differentiated, while allozymes revealed low levels of intrapopulational genetic variation. Within populations of a species, Bruederle and Fairbrothers (1986) found significant deviations from the Hardy-Weinberg expected heterozygosity and high positive values for Wright's fixation index suggesting high levels of inbreeding. The genetic structure of .C. .fuM! and .C. viridula is also similar to that reported for the

.C. crinita Lam. complex (Bruederle and Jensen 1991). Genetic diversity was

highest among populations within a taxon with relatively little variation found within populations. Low values for the number of alleles per polymorphic locus,

proportion of polymorphic loci, and Hardy-Weinberg expected heterozygosity

indicate that .C. .fuM! and .C. viridula are effecting a selfing or in-breeding

behavior (Bruederle and Jensen 1991). Genetic diversity was very low within

populations of _c. mendocenensis and .C. iYJlOdynama; and chromosome numbers

varied in both species but not within populations (Waterway 1990).

Genetics of the Carex brevior ~oup. Wahl (1940) collected chromosome

data from members of the .C. breyior group: .C. tenera n = 26, 27, and 28; .C. 32

normalis n = 34; _c. molesta n = 34; and _c. festucacea n = 34 + 3 trivalents.

The chromosomal races of _c. tenera exhibit some variation in form, with race n = 27 representing the larger, more robust extreme of the species. The trivalent in _c. festucacea usually consists of two medium and one small univalents (in

three out of 24 plates counted there were 35 bivalents and 1 univalents, the

smallest univalent separate from the other pair).

Summary of literature

Based on the literature, it is clear that members of the .C. brevior group

form an aneuploid series of morphologically similar, yet probably distinct taxa.

These species occur naturally in Iowa and are a subset of the .C. straminea

aggregate. A review of their taxonomic history clearly indicates a need to study

these species at the population level, at multiple sites, to determine which

characters (if any) can be used to reliably separate these taxa. If taxa exhibit low

genetic variability, as the literature suggests, then morphlogical variation occurring

within populations may be attributed to phenotypic plasticity. The significance of

taxonomic classifications can be determined using modem statistical techniques.

However, due to the nature of the group, any morphological differences will be

minute, requiring accurate measurement of morphological characters from a large

number of specimens in order to reduce sampling error. 33

CHAPTER2

ME1HODS

Herbarium Study

A preliminary survey of herbaria was undertaken to determine which carices of the section Ovales had previously been collected in Iowa. Specimens of the the Carex straminea aggregate were examined at the following herbaria:

University of Northern Iowa, Cedar Falls (ISTC), University of Iowa, Iowa City

(SUI), Iowa State University, Ames (ISC), and the Missouri Botanical Gardens,

St. Louis (Mo) (abbreviations taken from Holmgren et al. 1981). In order to compare taxonomic literature with historical collections and annotations, loans were obtained from the Gray Herbarium (GH), Bailey Hortorium (BH), Kansas

State University (KSU), Missouri Botanical Garden (MO), Philadelphia Academy of Natural Science (PH), and the New York Botanical Gardens (NY).

I visited PH and NY to view their extensive Carex collection and examine

specimens of the Carex straminea aggregate collected throughout their geographic

range. PH houses the exherbaria of Rev. G. H. E. Muhlenberg, who sent

numerous specimens to Willdenow (1805) for description, and that of Rev. Lewis

D. De Schweinitz, who with John Torrey, published one of the first critical

treatments of Carex in North America (Schweinitz 1824 1826). Most of the

specimens illustrated by H. C. Creutzburg for Mackenzie (1940) are found at PH.

Creutzburg's illustrations serve to fix Mackenzie's concepts of species to carefully

chosen specimens. In addition the Carex collection of Bayard Long, long time co­

worker of M. L Fernald is housed at PH. Both institutions contain several Carex 34 type specimens, especially PH. The Carex collection of K K Mackenzie, some

40,000 specimens, are housed at NY. Representative specimens were borrowed and compared directly to Iowa collections. No morphological data from these specimens were statistically analyzed.

Field Collections

During the growing seasons of 1989 and 1990, a comprehensive field

survey of mature fruiting plants was conducted in Iowa for the Carex brevior

group. Populations of these taxa were studied from the end of May through

August 1989. Two additional populations were visited in June 1990.

Potential study sites were obtained from labels of herbarium specimens

and from Iowa floristic studies (Eilers 1975; Roosa, Leoschke, and Eilers 1989).

Many former collection sites were revisited. However, most known habitats

previously occupied by the group had disappeared, e.g., ephemeral marsh borders,

savannas, woodland edges, disturbed grasslands, river raceways, sand dunes and

roadsides. Therefore, supplemental sites, with similar habitats, were obtained

from documents of the State of Iowa Preserves Advisory Board, Iowa Department

of Natural Resources, The Iowa Chapter of the Nature Conservancy and from

local county conservation boards. Over 100 sites were investigated throughout

Iowa, from which 22 populations were sampled from 15 sites, resulting in over

500 specimen collections.

At each study site, population boundaries for each taxon present were

delimited during a preliminary inspection of the area. Ecological attributes

within population boundaries were noted, including soil type, topography, and 35 associated species. The number of plants with fertile culms was estimated for each population. The plants to be sampled were selected according to predetermined numbers randomly generated by a Hewlett-Packard HP-llC

calculator. The calculator would generate a decimal value to the hundredths position which was then multiplied by the population size estimate to obtain a whole number. Plants were counted as I walked through the population along a prescribed route, always beginning at the point where I first discovered the population. Along the route, plants were counted from left to right then right to left in about a 5 meter strip. As the count proceeded specimens assigned a predetermined number were collected as a random sample. For statistical

analysis, up to 30 plants with mature fruit were sampled from each population.

A sample was defined as those portions of a plant collected for character analysis

later in the laboratory. A plant (genet) was defined as all the culms (ramets)

generated from a common rootstock or short rhizome (Bernard 1990). Within

the plant, there can be morphological variation among individual ramets. This variation appears mainly among characters correlated with culm size and

developmental stage. In an effort to standardize the selection of vegetatative and

fertile culms from which characters were measured, I selected the tallest, most

robust culm of each plant, provided it had all the necessary structures.

Occasionally there were other ramets with unusual features. These were

measured separately as form B of the same sample. Unidentified specimens were

classified as unknowns and treated separately until their taxonomic status could

be ascertained. Separate groupings of plants within a larger site were considered

together as one population. 36

Laboratozy Evaluation

I measured 44 morphological characters (31 quantitative and 13 qualitative) from each dried specimen (Table 3). There were 32 characters measured of the infructescence, perigynium, and achene. The remaining 12 characters were obtained from vegetative and fertile culms. An additional 12 ratios of characters were calculated by the computer for each specimen. Features of the culm and infructescence were measured to the nearest em or mm as appropriate. Smaller reproductive structures were measured with an optical reticule (12 lines per mm) in a dissecting scope. When evaluating qualitative

characters, separate states were assigned a numerical value for statistical analyses.

Statistical Analyses

Morphological data were analyzed using univariate and multivariate

statistical procedures (SAS Institute Inc. 1988). These tested the hypothesis that

taxa within the Carex brevior group are morphologically distinct and identified

useful taxonomic characters. All samples were treated as preclassified

Operational Taxonomic Units (OTUs).

Univariate Analyses

The variables of each taxon were analyzed by the SAS Univariate

Procedure to obtain means, standard deviations, variances and frequency

distributions. Frequency distributions can indicate if further segregations within

taxa are needed. Deviations from a normal frequency distribution, such as a bi­

modal histogram, may indicate significant differences between populations within

a taxon. For each histogram, the SAS Univariate Procedure lists specimens with 37 the 5 smallest and 5 largest character values of the distribution. This is useful for recognizing specimens which may be misclassified or express character values intermediate with other taxa. The SAS t-test Procedure was performed on paired combinations of taxa in order to identify significant differences (l2 < 0.0001) between variable means.

Table 3. Suite of characters used in scoring specimens for analysis. Vegetative characters were excluded from multivariate analysis. Ratios were derived from measured attributes of characters for each specimen (see Figures 1, 2, and 3). Fertile Culm Characters: 1 Number of leaves 2 Culm width at widest point (mm). 3 Extended leaf height (em). 4 Culm height (em). 5 Height to upper leaf base (em). 6 Culm height to upper leaf base/ culm height (ratio 5/4). 7 Upper leaf length (em). 8 Upper leaf width (mm). Vegetative Culm Characters: 9 Number of leaves. 10 Culm width (mm). 11 Leaves extended height (em). 12 Culm height to upper sheath apex (em). Inflorescence Characters: 13 Inflorescence shape: {1) 1 inear {2) oblong (3) ovoid (4) globose 14 Inflorescence length (mm). 15 Inflorescence width at widest point (mm). 16 Inflorescence width/length (ratio 15/14). (table 3 continues) 38

Spike Characters (lnflorescent Units)(Reznicek 1990): 17 Number of spikes. 18 Spike arrangement: (1) closely aggregated (2) loosely aggregated to spreading (3) moniliform (spikes approximate) (4) strongly moniliform. 19 Spike length (mm). 20 Spike width at widest point (mm). 21 Spike width/length (ratio 20/19). 22 Spike shape: (1) subglobose (4) obovoid (2) globose (5) oblong (3) ovoid (6) turbinate-obconical. 23 Spike apex shape: (1) truncate (4) rounded (2) blunt (5) pointed. (3) tapering 24 Terminal spike base shape: (1) truncate (4) short clavate (2) rounded (5) clavate. (3) tapering 25 lateral spike base shape (See 24). 26 Cycles of male florets on terminal spike (Number of revolutions around spike). 27 Cycles of female florets on terminal spike (Number of revolutions around spike including apical floret). 28 Cycles of male florets on lateral spike (Number of revolutions around spike). 29 Cycles of female florets on lateral spike (Number of revolutions around spike including apical floret). 30 Terminal male cycles/terminal female cycles (ratio 26/27). 31 lateral male cycles/lateral female cycles (ratio 28/29). Perigynium Scale Characters: 32 Scale length (mm). 33 Scale width (mm). 34 Scale lengthjperigynia length (ratio 32/36). 35 Scale widthjperigynia width (ratio 33/37). (table 3 continues) 39

Perigynium Characters: 36 Perigynia length (mm}. 37 Perigynia width (mm}. 38 Perigynia length from base to widest point {mm}. 39 Perigynia width/length {ratio 37/36}. 40 Perigynia body shape width/length (Figure 1}: (1} narrowly elliptic (1:3} (2} elliptic (1:2} (3} elliptic (2:3) (4} widely elliptic (5:6) (5} circular (1:1) (6} subrotund (6:5) 41 Perigynia length to widest pointjperigynia length (ratio 38/36}. 42 Position of widest point on perigynium: (1) mid-point (2} various intermediate positions (3} lower third 43 Beak length {apex of achene to tip} {mm). 44 Beak length/perigynia length (ratio 43/36). 45 Beak shape (from perigynium mid-point to beak apex}: (1} tapered (2} slightly constricted (3} abruptly constricted 46 Number of ventral nerves on perigynium over achene. 47 Appearance of ventral nerves: (1} strongly elevated above periynium tissue (2} finely imbeded in perigynium tissue (3} barely visible (4} absent 48 Number of dorsal nerves over achene on perigynia. 49 Perigynia dorsal nerve quality (See No. 47). Achene Characters: 50 Achene length (mm). 51 Achene width (mm). 52 Achene width/length (ratio 51/50). 53 Achene shape width/length (see 40}. 54 Achene position of widest point (see 42}. 55 Achene lengthjperigynia length (ratio 50/36). 56 Achene width/perigynia width (ratio 51/37}. 40

1 2 3 4 5 6 1:3 1:2 2:3 5:6 1: 1 6:5

1 000000

2 ~ooooo

3 ~00000

Figure 1. Symmetric plane diagrams adapted from Radford et al. (1974). Horizontal numbers correspond to character states given in Table 3: numbers 40 and 53. Columnar numbers correspond to character states given in Table 3: numbers 42 and 54.

Multivariate Analysis

The validity of taxonomic classifications within the Carex brevior group can be tested using linear discriminant functions. These are F -statistics based on the generalized squared distances between taxon means or (cluster centroids when specimens are plotted in n-dirnensional space). In general, multivariate analysis enumerates the various states of a character along a vector. Specimens are plotted at a specific point along this vector depending on the magnitude of the character state they express. The vector's length is a measure of the magnitude of the total variation expressed by all the specimens displaying that character.

The set of character vectors which separates the most specimens becomes the first canonical axis. A canonical axis is a suite of correlated character vectors 41

.Qn:.g tenera Dewey Carex tenera var. echinodes (Fernald) Wiegand

a c a b

Carex festucacea Schkuhr ex Willdenow Carex normalis (Dewey) Mackenzie )\ ;~\ 0

c a b d a b c d

Carex brevior (Dewey) Mackenzie Carex molesta Mackenzie ex Bright

a b c d a b c d

Figure 2. Perigynia of the Carex brevior group (redrawn from Mackenzie (1940) except for ..C. tenera which is redrawn from Boott (1862). (a) Dorsal surface of perigynium, (b) ventral surface of perigynium, (c) sub tending perigynium scale, and (d) achene. 42

INFLORESCENCE SHAPE and SPIKE ARRANGEMENT

Infl. shape globose oblong to linear Spike arr. linear closely loosely strongly aggregate aggregate t moniliform spreading

.8.1h f. 7th f. CYCLES OF MALE AND 6th f. ~ f\ L\ FEMALE FLORETS 5thf. ''" - (~. brevior) 4th f. \'::(.. j~ 3rd f. - setaceous tip 2nd f. ~' \;Yf.. U of 1ateraJpi ke 1st 5th male c. 4th Iii. '»~~ :0~l-i: 3rd m.

bract

SPIKE SHAPES

globose spike ovoid spike oblong spike round base tapered base clavate base

..)

Figure 3. Features of the inflorescence and spike (adapted from Mackenzie (1940). 43 with a parallel direction. Characters correlated to the first canonical axis explain the largest proportion of the total variation of all the characters expressed by the dataset. Eigenvalues represent the proportion of the total variation explained by a particular canonical axis.

Each character is assigned a canonical coefficient which is the proportion of the character's variation correlated to a canonical axis. A canonical value for each character is determined by multiplying a character's canonical coeficient by the magnitude of the character state expressed by a specimen. A specimen is plotted along a canonical axis by summation of all its canonical values. When specimens exhibit several discriminating characters, they can be plotted along several canonical axes--each with it's own combination of correlated characters.

Canonical axes were orthogonally arranged. This results in a multidimensional ordination by which relationships between specimens are illustrated in n-dimensional space. In canonical plots, close proximity among a set of specimens suggests a relationship, such as members of the same taxon. This is illustrated in canonical plots as a cluster of specimens. The relative distance between cluster centroids compared to the average distance from the centroid to each of the plotted specimens within the cluster, can be used as a measure of significance to test whether the total variation between taxa is greater than the total variation within each taxon.

Discriminant Function Analysis (DFA) is a multivariate procedure used primarily to classify OTUs into two or more known classes on the basis of one or more variables. This contrasts with Principle Component Analysis which assumes 44 no prior class membership because its purpose is to construct a classification scheme. DFA is able to distinguish taxa which closely resemble one another by analyzing variables together even though individual variables overlap considerably between taxa. DFA has two important uses: 1) it can determine if taxonomic classifications are significantly different, and 2) it can evaluate the contribution each character makes in distinguishing taxa (Sneath and Sokal 1973). Stepwise

DFA lists characters with the most discriminating power according to the amount of overall variance explained by each character. DFA and Stepwise DFA was performed by SAS DISCRIM procedure. SAS DISCRIM Posterior Error-rate

Estimates evaluates the probability that a particular OTU was misclassified.

SAS Canonical Analysis is a DFA procedure which summarizes between­ class variation and is capable of plotting specimens on canonical axes. Each canonical plot was illustrated twice in order to depict relationships among specimens according to taxonomic classification and collection site. Specimens occupying intermediate positions between group clusters on cannonical plots were re-examined. Posterior classification of unknown specimens were made when such OTUs fell within a taxon cluster. 45

CHAPTER 3

RESULTS

Most of the field study was accomplished during the summer of 1989 and completed in June 1990 when populations of Carex festucacea and _c. molesta were sampled. Populations of the .C. brevi or group (Table 1) were sampled at 15 sites throughout Iowa (Table 4). Eight of these were sympatric sites with more than one taxon present (Table 5). The taxa were found in discrete habitats, albeit some were microsites with small environmental differences. There were

450 randomly collected specimens measured for univariate analysis. However, 82 specimens were excluded from multivaritate analysis because of missing values for one or more variables. Sample sites, population sample size, total sample size for each taxon (N), and sampling dates are provided in Table 5. Field observations coupled with herbarium study justify listing _c. tenera var. echinodes as a new taxon for Iowa.

Field Observations

Growth Habit of Iowa Ovales

Iowa carices of the section Ovales, (Table 2) exhibit a phalanx growth form (Bernard 1990). Individual plants (genets) reproduce vegetatively from short rhizomes becoming caespitose with numerous vegetative and fertile culms

(ramets). At anthesis, genets usually have well developed vegetative culms with nodes and vascular tissue. Within populations, vegetative culms of all developmental stages were present, ranging from those in the protective prophyll

(a coleoptile-like sheath which protects the meristem and leaves while 46

Table 4. Identification codes, site names and county location. CODE SITE NAME COUNTY LOCATION BSM Big Sand Mound Nature Preserve Lousia CHSP Cedar Hills Sand Prairie Black Hawk FA Falls Access Wildlife Area Black Hawk GWSP Geo. Wythe State Park Black Hawk HP Hayden Prairie State Preserve Howard TFSP Lake of Three Fires State Park Taylor MCP Martin County Park Cherokee OTC Orono Township Cemetary Muscatine PKSP Palisades-Kepler State Park Linn PMP Private Mesic Prairie Black Hawk RTP Rolling Thunder Prairie Warren SRCP Split Rock County Park Chickasaw SCP Starr's Cave Preserve Des Moines VLSP Viking Lake State Park Montogomery WP Williams Prairie TNC Preserve Johnson

they develop from the bud of the rhizome) to fully developed ramets with internodes. Some appeared to be pseudoculms or false stems made up of a series of overlapping leaf sheaths. Pseudoculms are recognized by "leaf bases borne very close together, as a rosette, on a tightly compacted stem without discernable internodes (Reznicek and Catling 1986)." These "pseudoculms" appear to be a developmental stage and had no taxonomic value within the _c. brevior group.

The meristems of late-growing vegetative shoots are capable of overwintering above ground, developing into fertile culms the following spring.

Iowa Ovales carices exhibit sympodial growth where culms develop from apical meristems of rhizomes. The rhizome forms lateral buds which may develop on any side of the developing culm. These lateral buds may produce culms and may eventually form into rhizome branches (Figure 4). The number of 47

1 2 Table 5. Taxon, site code, canonoical plot site code , sample size , other species 3 presence (mixed or sympatric populations) , and date collected. TAXON SITE CANON. SAMPLE MIXED DATE CODE SITE SIZE POP. COLLECTED

Carex brevior BSM 0 32 * 06-13/89 SCP m 2 06-15/89 CHSP 1 29 06-17/89 GWSP ; 20 06-22/89 PKSP h 10 * 06-26/89 TFSP g 19 * 06-27/89 RTP d 11 07-03/89 WP b 3 * 07-09-89 Total N 126 Carex festucacea BSM a 26 * 06-15L90 Carex molesta OTC n 25 * 06-14/89 SCP m 31 06-15/89 FA j 1 06-22/89 PKSP h 16 * 06-26/89 TFSP g 4 * 06-27/89 WP b 13 * 07-09/89 BSM 0 2 * 06-15/89 BSM a 28 * 06-15/90 Total N 120 Carex normalis SRCP p 2 * 06-09/89 SCP m 2 * 06-15/89 HP k 2 * 06-20/89 VLSP f 1 06-29/89 PMP c 22 07-07/89 Total N 29 Carex tenera SRCP p 25 * 06-09/89 HP k 26 * 06-20/89 Total N 51 Carex tenera var. echinodes MCP e 16 06-30/89 1See Figures 4-6: canonical plots of specimens by site. 2 A total of 450 specimens were collected of which 368 were statistically analyzed. 3(*) indicates sympatric site. 48 ramets per genet were observed to vary from 1 to 160 culms. The circumferences of genets measured at the base ranged from 3.5 em for a solitary ramet, to 110 em for 160 ramets. Diagrams of some observed growth patterns are given in

Figure 4.

I I + I --1--H t +

A B c D E Figure 4. Aerial views of rhizome growth patterns depicting locations of meristems and ramets along rhizome axes. (A) Initial culm developed from seed. (B) Rhizome elongating in 2 directions with 2 apical meristems and 6 culms. (C) Rhizome elongating in one direction with 1 apical meristem and 15 culms. (D) Rhizome appendages elongating in several directions at ends of 4 branches (rhizome has many meristems and culms at the ends, but is barren at center). (E) Fragmented rhizome branches elongating in one to many directions away from the origin.

In grassland habitats, .C. brevior and .C. bicknellii exhibited directional rhizome growth. These plants appeared to move along the ground in interstitial zones between clumps of grasses, forming new shoots at the growing end and dying at the other end. Often a rhizome would be devoid of shoots for much of 49 its length, having cu1ms only near the apical meristem. Clusters of culms, appearing to be individual genets separated by several centimeters, would sometimes be connected underground by rhizome branches. In other cases, decay would disintegrate the connecting rhizome leaving individual clusters of ramets scattered throughout an area of two square meters. Several clumps of _c. bicknellii ramets were seen at Hayden Prairie in a fairy-ring pattern appearing to originated from a central point. A plant was observed which aborted apical growth of the rhizome when it collided with the roots of Andropogon gerardii.

One of the lateral buds then resumed growth, resulting in a rhizome with a 90 degree bend. Generally, the rhizome produces a fibrous root mass (rhizosphere) which forms a thick mat below the soil surface. I found rhizosphere diameters about 3 times that of the genet base (10-20 ramets) in one population of _c. brevior growing in disturbed sandy soil. The fibrous root mass of _c. brevior provided sites for the establishment of other species. I observed seedlings of Poa pratense and forbs growing in the fibrous roots while none were observed in the soil composed of eolian sand alone. The rhizosperes of larger genets of _c. molesta (ramets > 100) occupied sub-surface areas up to a meter in diameter on mud flats along a creek.

Ecological Characteristics

Iowa Ovales are opportunistic species that produce several fertile culms.

They quickly dominate disturbed habitats such as marsh drawdowns, shifting sand,

sediment deposits and old fields. Ovales carices are weak perennials, and are

often replaced in many habitats by more aggresive species with stronger 50 rhizomatous growth. For example, without continuous disturbance, .,C. molesta would be crowded out of moist, organic soils by Phalaris arundinacea L In stable habitats, such as prairie grasslands or wet sedge-meadows, Ovales were found in microsites characterized by localized environmental stress or in ecotonal zones, e.g., woodland edges or interfaces between dry uplands and low wetlands.

Field observations of Carex brevior. At Big Sand Mound (BSM), a region of drifting sand along the Mississippi River, scattered genets of .C. breyior were found in swales, slope bases and slight depressions. Most individuals of the population were found along woodland edges. This distribution seems related to increased availablity of soil moisture and reduced competition. At the Cedar

Hills Sand Prairie Preserve ( CHSP}, a few individuals of .,C. brevi or were found persisting in an ecotonal zone between prairie grasses and tussocks of .,C. stricta.

However, a larger number of more vigorous plants were found in an old field of eolian sand currently reverting back to prairie .

.,C. brevior was also found in old pastures and prairies on loamy soil and on riparian sand bars. At BSM, .,C. brevior was associated with Ulmus pumila,

Calmovilfa lon~folia. Era~ostis trichodes. Koleria macrantha. Andropogon gerardii, Carex muhlenber~a. .s.t.ip,a spp., Asclepias amplexicaulis, Helianthus annuus, Plantago aristata, Tephrosia vir~niana, Lithospermum canescens,

Tradescantia ohioensis, Amorpha canescens, Mirabilis nyctaginea, and

Chenopodium spp.

Field observations of Carex molesta. .,C. molesta was often found near .C. brevior at many sites, but in different microhabitats. .,C. molesta is larger and 51 more robust, tending to favor moist, organic soils while .C. brevior favors dryer, sandier soils. During sampling, this difference would not be immediately detected because of specimens with intermediate characteristics. The best example of this occured at Palisades-Kepler State Park (PKSP) on an old river channel transecting a meander. There were natural sand levies at both ends, creating a linear, stagnant pool which had been receding. After the initial survey, I began sampling .C. brevior on the sandy soil on top of the levy. As I sampled down the

12 foot slope toward the water, I collected samples which displayed intermediate character states. These specimens were difficult to classify. Typical forms of .C.

molesta were found on slope bases in wet soil composed of organic silts and sand.

Both species were also sympatric at William's Prairie Preserve (WP) which has a

swale topography. .C. brevior was found on small ridges and .C. molesta was

found in the occasionally flooded swales.

Carex molesta was also sympatric with .C. festucacea at BSM, where large

genets of both taxa were intermingled on clayey sub-soil exposed when a 6 meter

high levy was constructed along the Mississippi river. They and Ambrosia

artemisiifolia were the dominant herbaceous cover. Even though ramets of both

taxa were intertwined together, they were easily distinguished by features of the

perigynia and inflorescence .

.C. molesta was also found in the partial shade of alluvial and upland

woods, wet-mesic prairies, and roadside ditches. Associated species at PKSP were

Phalaris arundinacea. Elymus virginicus, .C. stipata. .C. cristatella. .C. tribuloides, C.

annectens, .C. breyior, Solidaio iiiantea. Ru

Field Observations of Carex festucacea. There was only one population of

.C. festucacea sampled (BSM). A solitary specimen was collected in a

depauperate upland prairie at Gull Point State Park ( GPSP) in Dickinson County,

lA. This specimen appears to be a disjunct population because all the historical

collections were made in eastern Iowa. In addition, Wheeler and Ownbey (1984)

report the only Minnesota collection site of ,C. festucacea is north of Dickinson

County, lA.

At BSM, ,C. festucacea was sympatric with both ,C. molesta and .C. brevior.

Specimen # F:lOla was collected with ,C. molesta in partial shade at the base of

large sand dune. Its perigynia resembles ,C. molesta in being slightly longer and

lanceolate shaped; however, its spikes are obovate and the inflorescence is

moniliform. Most of the specimens of .C. festucacea at BSM were found on

exposed clay subsoil. The specimens at GPSP was collected on an upland prairie

at the edge of a trail which cut into a slope.

Field Observations of Carex tenera. Scattered plants of ,C. tenera were

found in wet-meadows and grasslands. Genets were found growing among several

species at Hayden Prairie Preserve (HP) and Split Rock County Park (SRCP).

Genets usually had a small number of ramets (N ...:5.. 20), but some were

occasionally found with up to 60 ramets. The slender culms were weak and

frequently nodding. ,C. tenera perferred open areas in moist organic soils

bordering wetter soils. Disturbances to the habitat favored C. tenera, e.g., at

SRCP most of the plants were found in an area drained by a deeply cut road

ditch. Other forms of disturbance to the natural community were evident. 53

There were an estimated 350 genets of ..C. tenera in about 3 acres at SRCP. They were in wet-mesic meadows, bordering areas dominated by ..C. stricta, ..C. hystricina and ..C. suberecta. ..C. tenera was associated with the following forbs: Silphium perfoliatum, Geum triflorum, Zizia ~ Saxifraga pensylvanica, Veronicatrum viq~inicum and C.ypripedium candidum. At Hayden prairie, ..C. tenera was found in a narrow ecotonal strip between borders of prairie uplands and wet sedge-meadows bordering a prairie creek. Occasionally it was found at higher elevations in depressions and swales amid the upland prairie.

Several ramets of _c. tenera were seen in fire-breaks mowed the previous year. This form of disturbance greatly favored _c. tenera over more competitive species.

Field Observations of Carex normalis. Small numbers of C. normalis plants were infrequently observed in open oak woods throughout the state. The plant closely resembles ..C. sparganioides which is more common in the same woodland habitats. This species has androgynous spikes with male florets at the

apex, but its vegetative features are nearly indistinguishable from ..C. normalis.

The spreading perigynia of both species are loosely held by the inferior scales and

are easily shattered from the spike. The largest population of _c. normalis was

sampled at Viking Lake State Park (VLSP) where they occupied a transitional zone between the Bromis inermis bordering the lake, and the densely shaded

understory of an oak forest. The plants were scattered among a brier patch of

~ missouriensis and Rubus alle~heniensis. Unfortunately, only one of these

plants could be analyzed by multivariate methods because of missing perigynia.

However, a second population was measured in the field before the perigynia 54 shattered. These were found at a six acre private mesic prairie (PMP) near

Cedar Falls. About 30 genets of .C. normalis occupied a 3 meter by 1 meter strip of prairie which received sediments from an adjacent cultivated field. Plants were also found in dirt piles and flood plains. Two plants of .C. normalis were collected at SRCP in drained loamy soil beneath an aspen thicket bordering a sedge-meadow where .C. tenera was found.

At Viking Lake State Park (VLSP), .C. normalis was associated with Carya

.QYata, Quercus borealis, ..Q. macrocarpa, Ribes missiourense, Zanthoxylum

americanum, Rubus alleiheniensis, Toxicodendron rydberiii, Desmodium ilutinosum, .fQa pretensis, Elymus villosa, and C. sparianioides.

Field Observations of Carex tenera var. echinodes. The taxonomic status of .C. tenera var. echinodes has been uncertain since its original description

(Fernald 1902). The vegetative features are similar to .C. tenera with narrow leaves and culms. The spikes are arranged in a bead-like pattern (moniliform) along the rachis which is thin and flexuous. This gives the inflorescence a nodding or zig-zag appearance. The characteristics of the perigynia and spikes closely resemble those of .C. normalis except the perigynia are slightly larger. I found only one population of .C. tenera var. echinodes in wet woods. Most of the plants were located in a roadside where water seeped into the ditch at Martin

County Park (MCP). A few plants were scattered in a densely forested ravine along an intermittant stream. Such a habitat is very different from either the open meadows of .C. tenera or the well-drained savanna of .C. normalis. Associate species at MCP were Caxya cordiformis, Ulmus 1ll1ml, .Iiliil americana, 55

~ ne~ndo, Prunus americana. Jun~lans .nJirn, Ostzya yir~ana., San~inea

canadensis. Poly~onatum biflora, Smilicina racemosa, Viburnum rafinesq_ueii,

Triosteum perfoliatum. Rhamnus cathardica, Osmorhiza lon~stylis, Hydrophillum

vir~nianum, and .C. blanda. Statistical Analyses

Univariate Analyses

Univariate analysis was performed on the 43 measured characters and 13

computer derived character ratios (Table 3) to obtain means, standard deviation

and minimum-maximum values (Tables 6). Comparative box plots depicting

means, modes, interquartile range (an interquartile range is the distance betweent

the 25th and the 75th sample percentiles), and minimum-maximum ranges are

given for important taxonomic characters (Figure 5). Box plots represent

histograms. Characters with normal distributions have means equal to the modes,

equal distances to interquartile boundaries, and equal length of the tails. A

character with a leptokurtic distribution has a narrow interquartile range where a

disproportion of the specimens have character states near the mean. A character

with a skewed distribution to the right has a mode separate and to the right of

the mean; and the left tail is long, extending the lower boundary further away

from the mean than the upper boundary.

Test of normality. Normal frequency distributions are necessary for

multivariate analysis by parametric methods. Normal distributions were obtained

for most characters of .C. brevior (N = 147) and _c. molesta (N = 131) 56

Table 6. Character description, taxon, sample size (N), mean, standard deviation (SD), mininium value and maximum value. Taxon H Mean so Minimum Maximum -

1. Fertile culm number of leaves~

~- breyior 147 3.24 0.72 2.00 5.00 ~- tenera var. echinodes 29 3.59 0.78 3.00 6.00 ~- festticacea 31 3.03 0.55 2.00 4.00 ~- molesta 131 3.54 0.83 2.00 6.00 ~- normalis 52 3.98 0.70 2.00 6.00 ~- tenera 58 3.50 0.57 2.00 5.00 2. Fertile culm width at widest point (mrn).

~- breyior 147 2.10 0.42 1.00 3.00 ~- tenera var. echinodes 29 1.88 0.29 1.50 2.50 ~. festycacea 33 2.06 0.24 1.50 3.00 ~. molesta 131 2.17 0.48 1.00 3.50 ~. normalis 52 3.11 0.39 2.50 4.00 ~- tenera 58 1.86 0.31 1.00 2.50 3. Fertile culm extended leaf height (em).

~. breyior 141 35.84 8.94 18.00 65.00 ~. tenera var. echinodes 27 55.22 7.61 44.00 71.00 ~. festycacea 31 44.06 10.75 25.00 77.00 ~- molesta 128 51.63 10.72 31.00 76.00 ~. normalis 51 69.69 11.64 39.00 104.00 ~- tenera 58 48.72 7.82 29.00 77.00 4. Fertile culm height (em).

~. brevior 147 63.26 19.01 29.00 116.00 ~. tenera var. echinodes 29 80.00 11.06 56.00 99.00 ~. festycacea 33 84.15 14.84 56.00 117.00 ~. molesta 131 78.94 13.50 36.00 112.00 ~- normal is 52 104.42 15.20 61.00 142.00 ~. tenera 58 59.28 8.53 41.00 76.00 (table continues) 57

Taxon t! Mean so Minimum Maximum

5. Culm height to upper leaf base (em).

~. brevior 141 18.49 6.08 7.00 38.00 ~. tenera var. echinodes 27 24.93 5.00 17.00 37.00 ~. festycacea 31 22.97 7.14 11.00 41.00 ~. molesta 128 27.98 7.56 13.00 50.00 ~. normalis 51 32.98 6.61 0.00 47.00 ~. tenera 58 21.67 4.74 9.00 36.00 6. Ratio: Fertile culm height to upper leaf base/fertile culm height.

~. brevior 141 0.30 0.08 0.16 0.81 ~. tenera var. echinodes 27 0.31 0.05 0.23 0.42 ~. festucacea 31 0.27 0.06 0.20 0.46 ~. molesta 128 0.36 0.08 0.21 0.56 ~. normalis 51 0.32 0.07 0.00 0.52 ~. tenera 58 0.37 0.06 0.22 0.52 7. Fertile culm upper leaf length (em).

~. brevior 142 17.30 4.40 7.00 31.00 ~. tenera var. echinodes 27 30.30 4.74 23.00 46.00 ~. festycacea 31 21.10 4.44 12.00 36.00 ~. molesta 128 23.65 5.05 14.00 38.00 ~. normal is 51 36.71 8.51 27.00 64.00 ~. tenera 58 27.05 4.77 15.00 41.00 8. Fertile culm upper leaf width (nm).

~. breyior 146 2.46 0.48 1.50 3.50 ~. tenera var. echinodes 29 2.41 0.46 1.50 3.00 ~. festycacea 28 2.84 0.31 2.00 3.50 ~. molesta 130 2.79 0.47 1.50 4.00 ~. normalis 52 3.82 0.44 3.00 5.00 ~. tenera 58 2.11 0.38 1.00 3.00 (table continues) 58

Taxon t! Mean so Minimum Maximum

9. Vegetative culm number of leaves.

~. brevior 66 7.35 1.58 3.00 12.00 ~. tenera var. echinodes 29 8.24 1.15 7.00 10.00 ~. festycacea 9 9.11 1.36 7.00 12.00 ~. molesta 55 8.22 1.97 3.00 13.00 ~. normalis 32 7.44 1.19 5.00 10.00 ~. tenera 33 4.97 1.26 0.00 7.00 10. Vegetative culm width {mm).

~. brevjor 67 1.86 0.55 1.00 3.00 ~. tenera var. echjnodes 29 1. 74 0.41 1.00 2.50 ~. festycacea 9 1.61 0.49 1.00 2.50 ~. molesta 55 1.87 0.42 1.00 3.00 ~. normal is 32 2.78 0.52 1.50 4.00 ~. tenera 33 1.44 0.46 0.00 2.00 11. Vegetative culm extended leaves height {em).

~. brevior 67 34.76 8.81 20.00 67.00 ~. tenera var. echinodes 29 56.28 6.84 42.00 66.00 ~. festycacea 9 55.22 11.29 41.00 79.00 ~. molesta 55 50.31 14.48 11.00 82.00 ~. normal is 32 63.38 18.68 22.00 110.00 ~. tenera 33 38.88 11.25 0.00 60.00 12. Vegetative culm height to upper sheath apex {em).

~. breyior 67 14.54 5.21 5.00 33.00 ~. tenera var. echinodes 29 23.97 4.91 14.00 33.00 ~. festycacea 9 27.22 7.17 19.00 42.00 ~. molesta 55 24.49 8.91 7.00 47.00 ~. normalis 32 23.66 8.83 10.00 42.00 ~. tenera 33 13.39 4.95 0.00 21.00 {table continues) 59

Taxon H Mean so Minimum Maximum

13. Inflorescence shape (See Table EE for quality states). t_. breyior 147 1.61 0.81 1.00 3.00 t_. tenera var. echinodes 29 1.00 0.00 1.00 1.00 t_. festucacea 33 1.53 0.62 1.00 3.00 t_. molesta 131 2.58 0.87 1.00 4.00 t_. normal is 52 1.92 0.96 1.00 3.00 t.. tenera 58 1.14 0,40 1.00 3.00 14. Inflorescence length {mm). t_. breyior 147 29.18 7.60 13.00 52.00 t.. tenera var. echinodes 29 39.21 7.33 24.00 53.00 t_. festycacea 33 41.67 8.20 27.00 58.00 t_. molesta 131 21.00 3.59 13.00 34.00 t_. normal is 52 27.60 5.14 16.00 45.00 t_. tenera 58 36.34 7.13 22.00 57.00 15. Inflorescence width (mrn). t_. breyior 145 10.30 2.31 5.00 18.00 t.. tenera var. echinodes 24 9.25 I. 73 5.00 12.00 t.. festycacea 33 10.03 2.05 5.00 15.00 t_. molesta 131 12.26 I. 75 7.00 16.00 t.. normal is 39 9.74 1.94 6.00 16.00 t.. tenera 58 7.91 1.61 5.00 13.00 16. Ratio inflorescence width to length. t.. brev1or 145 0.37 0.11 0.15 0.77 t_. tenera var. echinodes 24 0.24 0.07 0.13 0.37 t. festycacea 33 0.25 0.08 0.12 0.44 t. molesta 131 0.60 0.11 0.32 0.92 t.. normal is 39 0.34 0.08 0.19 0.55 t.. tenera 58 0.22 0.06 0.10 0.43 (table continues) 60

Taxon H Mean so Minimum Maximum

17. Spike number. k· brevior 146 4.18 0.80 3.00 6.00 k· tenera var. echinodes 29 5.79 0.86 5.00 8.00 k· festycacea 33 6.06 1.27 4.00 9.00 k· molesta 131 4.15 0.96 2.00 9.00 k· normalis 52 7.21 1.30 4.00 10.00 k· tenera 58 4.98 0.98 3.00 7.00 18. Spike arrangement (See Table EE.for quality states). k· brevior 146 2.45 0.93 1.00 4.00 k· tenera var. echinodes 29 3.52 0.63 2.00 4.00 k· festycacea 33 2.76 0.85 1.00 4.00 k· molesta 131 1.39 0.57 1.00 3.50 ~. normalis 52 1. 72 0.61 1.00 4.00 k· tenera 58 3.48 0.63 2.00 4.00 19. Spike length (mm). k· breyior 146 12.15 2.12 7.00 17.00 k· tenera var. echinodes 19 10.26 1.33 8.00 13.00 k· festycacea 33 11.52 2.09 7.00 15.00 k· molesta 130 10.49 1.91 6.00 16.00 k· normalis 32 10.00 1.68 7.00 13.00 k· tenera 58 10.33 1.98 6.00 15.00

20. Spike width (mm).

~. breyior 145 6.48 0.94 4.00 12.00 k· tenera var. echinodes 19 6.37 0.96 4.00 8.00 ~. festycacea 33 5.82 1.04 4.00 10.00 k· molesta 130 7.50 1.02 5.00 12.00 k· nonnalis 32 5.72 0.92 4.00 7.00 k· tenera 58 5.43 0.75 4.00 7.00 (table continues) 61

Taxon ti Mean so Minimum Maximum

21. Ratio: Terminal spike width/length. t,. brevior 145 0.55 0.10 0.31 1.00 ~. tenera var. echinodes 19 0.63 0.10 0.36 0.80 ~. festycacea 33 0.52 0.11 0.38 0.77 ~. molesta 130 0.73 0.14 0.50 1.20 ~. normalis 32 0.58 0.13 0.42 1.00 ~. tenera 58 0.54 0.11 0.36 1.00 22. Spike shape (See Table EE for quality states}.

~. breyior 142 3.87 0.65 2.00 6.00 ~. tenera var. echinodes 19 4.63 0.96 4.00 6.00 ~. festycacea 33 3.76 0.66 1.00 5.00 ~. molesta 129 2.95 0.55 2.00 4.00 ~. normal h 31 3.68 0.60 2.00 4.00 ~. tenera 57 4.02 0.44 3.00 6.00 23. Spike apex shape (see Table 3 for quality states}.

~. breyior 138 3.43 0.87 1.00 5.00 ~. tenera var. echinodes 19 2.63 0.76 2.00 4.00 ~. festycacea 32 2.50 1.02 2.00 6.00 ~. molesta 129 3.69 0.50 1.40 4.00 ~. normal is 31 2.81 0.70 2.00 5.00 ~. tenera 57 3.37 0.86 2.00 5.00 24. Terminal spike base shape (see Table 3 for quality states}.

~. brevior 147 4.14 0.96 1.00 5.00 ~. tenera var. echinodes 29 4.72 0.59 3.00 5.00 ~. festycacea 33 4.33 0.82 3.00 5.00 ~. molesta 131 2.33 0.79 1.00 4.50 ~. normal is 52 3.36 1.12 1.00 5.00 ~. tenera 58 4.33 0.76 3.00 5.00 (table continues) 62

Taxon tf Mean so Minimum Maximum

25. Lateral spike base shape {see Table 3 for quality states). t,. brevior 146 3.25 0.98 1.00 5.00 t,. tenera var. echinodes 29 3.14 0.35 3.00 4.00 t,. festycacea 33 3.73 0.94 2.00 5.00 t,. molesta 131 1. 78 0.50 1.00 3.00 t,. normal is 51 2.11 1.02 1.00 5.00 t,. tenera 57 3.28 0.64 2.00 5.00 26. Terminal spike number of cycles of male florets. t,. brevior 147 5. 77 2.18 1.00 11.00 t,. tenera var. echinodes 29 4.38 1.15 2.00 7.00 t,. festycacea 33 9.70 4.06 0.00 17.00 t,. molesta 130 4.27 1.89 0.00 9.00 .t,. normalis 52 5.04 2.55 0.00 15.00 t,. tenera 58 5.24 1.94 2.00 11.00 27 . Terminal spike number of cycles of female florets.

.t,. brevior 147 11.57 2.74 6.00 24.00 t,. tenera var. echinodes 29 11.10 1.93 8.00 15.00 t,. festycacea 33 18.64 3.64 9.00 27.00 t,. molesta 130 15.96 3.04 9.00 24.00 .t,. normalis 52 15.31 2.40 10.00 22.00 .t,. tenera 58 10.24 3.52 5.00 16.00 28. Lateral spike number of cycles of male florets. t,. brevior 146 4.91 2.24 1.00 11.00 t,. tenera var. echinodes 29 2.79 0.98 1.00 4.00 .t,. festycacea 33 8.15 4.42 1.00 17.00 t,. molesta 130 2.87 1.22 0.00 7.00 .t,. norma 1is 51 3.33 1.62 1.00 9.00 .t,. tenera 57 3.75 1. 73 1.00 9.00 {table continues) 63

Taxon ti Mean so Minimum Maximum

29. Lateral spike number of cycles of female florets.

~. brevior 146 11.64 2.78 6.00 22.00 ~. tenera var. echinodes 28 10.07 2.07 6.00 14.00 ~. festycacea 33 20.36 3.39 12.00 26.00 ~. molesta 130 15.08 3.24 0.00 25.00 ~. normal is 51 15.08 2.42 10.00 21.00 ~. tenera 57 11.21 3.69 5.00 19.00

30. Ratio: terminal spike male cycles to female cycles.

~. brevior 147 0.53 0.25 0.07 1.14 ~. tenera var. echinodes 29 0.41 0.13 0.20 0.67 ~. festycacea 33 0.56 0.31 0.00 1.42 ~. molesta 130 0.27 0.12 0.00 0.56 ~. normal is 52 0.34 0.23 0.00 1.50 ~. tenera 58 0.55 0.23 0.15 1.00 31. Ratio: lateral spike male cycles/female cycles.

~. brevior 146 0.45 0.23 0.06 1.11 ~. tenera var. echinodes 28 0.29 0.11 0.10 0.67 ~. festycacea 33 0.43 0.29 0.05 1.33 ~. molesta 129 0.19 0.08 0.05 0.40 ~. normal j s 51 0.22 0.11 0.05 0.64 ~. tenera 57 0.36 0.20 0.12 1.00 32. Scale length (mrn).

~. brev1or 147 3.35 0.34 2.50 4.42 ~. tenera var. echjnodes 29 2.76 0.15 2.50 3.08 ~. festycacea 33 2.64 0.22 2.17 3.00 ~. molesta 131 3.00 0.26 2.33 3.50 ~. normalis 52 2.57 0.28 1.92 3.00 t,. tenera 58 2.78 0.20 2.33 3.33 (table continues) 64

Taxon tf Mean so Minimum Maximum

33. Scale width (mm). t,. breyior 147 1.29 0.17 0.92 1.83 t,. tenera var. echinodes 29 1.17 0.11 0.92 1.42 t,. festycacea 33 1.14 0.13 0.92 1.50 t,. molesta 131 1.17 0.13 0.83 1.50 t,. normal is 52 1.23 0.18 0.92 1. 75 t,. tenera 58 1.13 0.16 0.83 1. 50 34. Ratio: scale length/perigynium length. t. breyior 146 0.92 0.10 0.66 1.23 t,. tenera var. ech1nodes 29 0.66 0.05 0.57 0.80 t,. festycacea 33 0.88 0.10 0.70 1.09 t,. molesta 131 0.74 0.06 0.60 0.91 t,. normal is 52 0.75 0.08 0.53 1.03 t,. tenera 54 0.94 0.11 0.67 1.26

35. Ratio: scale width/perigynium width. t,. brevior 146 0.51 0.08 0.37 0.78 t,. tenera var. echinodes 29 0.68 0.08 0.55 0.83 t,. festycacea 33 0.64 0.10 0.42 0.85 t,. molesta 131 0.51 0.06 0.38 0.75 t,. normal is 52 0. 71 0.12 0.48 1.13 t,. tenera 54 0.69 0.13 0.45 1.00 36. Perigynium length (mm). t,. breyior 146 3.67 0.43 2.67 5.17 "-· tenera var. ecbinodes 29 4.18 0.31 3.67 4.75 t,. festycacea 33 3.04 0.33 2.50 3.92 t,. molesta 131 4.06 0.32 3.08 4.75 t,. normalis 52 3.42 0.37 2.75 4.08 t,. tenera 54 3.02 0.34 2.17 3.83 (table continues) 65

Taxon tf Mean so Minimum Maximum

37 . Perigynium width (mrn).

.t_. brevior 146 2.56 0.31 1.50 3.25 .t_. tenera var. echinodes 29 1. 74 0.14 1.50 2.08 .t_. festucacea 33 1.80 0.19 1.42 2.25 .t_. molesta 131 2.32 0.20 1. 75 2.92 .t.. normal is 52 1. 75 0.23 1.33 2.25 ~. tenera 54 1.65 0.20 1.00 2.00 38. Perigynium length from base to widest point (mm).

.t_. brevior 146 1.32 0.21 0.75 1. 75 .t_. tenera var. echinodes 29 1.11 0.19 0.83 1.50 .t_. festucacea 33 1.01 0.15 0.75 1.33 .t_. molesta 131 1.37 0.18 1.00 1.92 ~. normalis 52 0.98 0.16 0.58 1.25 ~. tenera 54 0.96 0.15 0.58 1.42 39. Ratio: perigynium width/perigynium length.

~. brevior 146 0.70 0.08 0.45 0.97 ~. tenera var. echinodes 29 0.42 0.04 0.35 0.53 .t_. festycacea 33 0.60 0.07 0.45 0.82 .t_. molesta 131 0.57 0.05 0.45 0.73 ~. normal is 52 0.51 0.06 0.38 0.70 ~. tenera 54 0.55 0.07 0.36 0. 71

40. Perigynium shape-- width/length (see Table 3 for quality states).

~. brevior 146 6.87 0.65 6.00 8.00 ~. tenera var. echinodes 29 4.00 0.00 4.00 4.00 ~. festycacea 33 6.55 0.62 5.00 8.00 ~. molesta 131 5.94 0.76 4.00 7.00 ~- normal is 52 5.35 1.43 3.00 8.00 ~. tenera 54 5.41 0.60 4.00 7.00 (table continues) 66

Taxon H Mean so Minimum Maximum

41. Ratio: perigynium length to widest pointjperigynium length.

~. breyior 146 0.36 0.04 0.24 0.49 ~. tenera var. echjnodes 29 0.26 0.03 0.21 0.32 ~. festucacea 33 0.33 0.03 0.26 0.41 ~. molesta 131 0.34 0.03 0.24 0.45 ~. normal js 52 0.29 0.03 0.21 0.34 ~. teneta 54 0.32 0.04 0.24 0.39 42. Perigynium shape-- position of widest point position (Table 3).

~. brey1or 146 1.45 0.86 1.00 4.00 ~. tenera var. echinodes 29 3.00 0.00 3.00 3.00 ~. festycacea 33 1.06 0.35 1.00 3.00 ~. molesta 131 1.71 0.97 1.00 4.00 ~. normal1s 52 2.08 1.01 1.00 3.00 ~. tenera 54 2.96 0.27 1.00 3.00 43. Beak length (mm) (as measured from apex of achene to beak apex).

~. breyior 146 1.71 0.33 1.08 2.75 ~. tenera var. echinodes 29 2.27 0.24 1.92 2.67 ~. festycacea 33 1.35 0.22 1.00 1.83 ~. molesta 131 2.16 0.26 1.50 3.00 ~. normal1s 52 1. 72 0.25 1.25 2.33 ~. tenera 54 1.52 0.22 0.92 1.92 44. Ratio: beak Jength/per1gynium length.

~. breyior 146 0.46 0.06 0.36 0.79 ~. tenera var. echinodes 29 0.54 0.04 0.46 0. 71 ~. festycacea 33 0.44 0.04 0.36 0.50 ~. molesta 131 0.53 0.04 0.36 0.65 ~. normalis 52 0.50 0.04 0.43 0.58 ~. tenera 54 0.50 0.06 0.39 0.71

(table continues) 67

Taxon .ti Mean so Minimum Maximum

45. Beak taper {see Table 3 for quality states).

~. brevior 146 2.64 0.53 1.00 3.00 ~. tenera var. echinodes 29 1.12 0.32 1.00 2.00 ~. festycacea 33 2.00 0.72 1.00 3.00 ~. molesta 131 1.59 0.45 1.00 3.00 ~. normal is 52 1.13 0.30 1.00 2.00 ~. tenera 54 1.55 0.84 1.00 4.00 46. Number of nerves on ventral face of perigynium over achene.

~. brevior 146 0.40 0.98 0.00 6.00 ~. tenera var. echinodes 29 3.45 1.15 1.00 5.00 ~. festycacea 33 3.18 1.57 0.00 6.00 ~. molesta 131 3.15 1.61 0.00 6.00 ~. normal is 52 4.38 1.40 0.00 8.00 ~. tenera 53 4.34 1.09 3.00 7.00 47. Ventral nerve appearance {see Table 3).

~. brevior 146 3.79 0.56 1.00 4.00 ~. tenera var. echinodes 29 1.26 0.54 1.00 3.00 ~. festycacea 33 2.53 1.01 1.00 4.00 ~. molesta 131 2.46 1.02 1.00 4.00 ~. normalis 52 1.66 0.83 1.00 4.00 ~. tenera 53 1. 74 0.74 1.00 3.00 48. Number of nerves on dorsal face of perigynium over achene.

~. brevior 146 2.99 1.46 0.00 7.00 ~. tenera var. echinodes 29 5.66 1.26 3.00 8.00 ~. festycacea 33 5.33 1.51 0.00 7.00 ~. molesta 131 5.02 1.23 2.00 8.00 ~. normali s 52 6.63 1.19 3.00 9.00 ~. tenera 53 6.94 0.97 5.00 9.00 {table continues) 68

Taxon tf Mean so Minimum Maximum

49. Dorsal nerve appearance.

~. brevior 146 2.27 0.96 1.00 4.00 ~. tenera var. echinodes 29 2.24 0.56 1.00 3.00 ~. festycacea 33 2.11 0.57 1.00 4.00 ~. molesta 131 2.06 0.62 1.00 3.50 ~. normalis 52 2.26 0.47 1.00 3.00 ~. tenera 53 2.37 0.55 1.00 3.50

50. Achene length (mm).

~. brevior 146 1. 75 0.12 1.42 2.00 ~. tenera var. echinodes 29 1.67 0.10 1.50 1.83 "-· festucacea 33 1.56 0.09 1.42 1. 75 ~. molesta 129 1.66 0.11 1.42 2.00 ~. normal is 52 1.45 0.12 1.17 1.67 ~. tenera 52 1.40 0.09 1.17 1.58 51. Achene width (mm).

~. brevior 146 1.41 0.12 1.00 1.67 ~. tenera var. echinodes 29 1.01 0.05 0.92 1.08 ~. festycacea 33 1.05 0.09 0.92 1.33 ~. molesta 129 1.16 0.08 0.92 1.42 ~. normalis 52 1.04 0.10 0.83 1.33 ~. tenera 52 1.03 0.07 0.92 1.17 52. Ratio: achene width/length.

~. brevior 146 0.81 0.06 0.65 1.00 ~. tenera var. echinodes 29 0.61 0.04 0.52 0.72 ~. festycacea 33 0.68 0.05 0.58 0.83 ~. molesta 129 0.70 0.05 0.58 0.89 ~. normal is 52 0.72 0.07 0.59 0.89 ~. tenera 52 0.74 0.06 0.61 0.87 (table continues) 69

Taxon tf Mean so Minimum Maximum

53. Achene shape-- width/length (see Table 3 for quality states).

,C.. brevior 145 6.27 0.69 4.00 8.00 ~. tenera var. echinodes 29 4.76 0.44 4.00 5.00 ~. festucacea 33 5.61 0.56 5.00 7.00 ~. molesta 128 5.52 0.52 5.00 7.00 ~. normalis 52 5.63 0.63 4.00 7.00 ~. tenera 52 5.21 0.89 3.00 6.00 54. Achene shape-- position of widest point (Table 3).

,C.. brevior 145 2.26 1.14 1.00 4.00 ~. tenera var. echinodes 29 2.59 0.82 1.00 3.00 ,C.. festycacea 33 1.30 0.73 1.00 3.00 ,C.. molesta 128 1.82 1.29 1.00 4.00 ,C.. normalis 52 2.10 1.09 1.00 4.00 ~. tenera 52 2.46 0.90 1.00 3.00 55. Ratio: achene lengthjperigynium length.

~. brevior 146 0.48 0.04 0.35 0.57 ~. tenera var. echinodes 29 0.40 0.03 0.32 0.45 ~. festycacea 33 0.52 0.05 0.44 0.67 ~. molesta 129 0.41 0.03 0.34 0.50 ~. normalis 52 0.43 0.03 0.36 0.50 ~. tenera 52 0.47 0.05 0.38 0.62 56. Ratio: achene width/perigynium width.

~. breyior 146 0.55 0.06 0.42 0.83 ~. tenera var. echinodes 29 0.58 0.05 0.48 0.67 ~. festycacea 33 0.59 0.04 0.48 0.67 ~. molesta 129 0.50 0.04 0.42 0.63 ~. normal is 52 0.60 0.06 0.50 0.75 ~. tenera 52 0.63 0.07 0.50 0.92 Character 2 Fertile culm width Character 8 Fertile culm upper leaf width 4 5 0 3.75 4.5 ___!_ __ 3.5 4 .1 ,.J., 3.25 3.5 3 3 2.5 .J .. 2.75 I . IH ·8·JJ, 'TIT''LIT' T 2.5 . . I . . . I 2 1- 1.5 ITI 0 2.25 T 2 H ·-·-----·-·1-·-----·-·H ·-·-----·-· ------·------·------·------·------·------·------0 1. 75 8 E F H N T 1.5 Itt. I 1. 25 B E F H N T Character 13 Inflorescence shape 4 3.75 Character 7 Fertile culm upper leaf length 3.5 65 3.25 60 3 ,_.__,__.,. _____ . 55 8 2. 75 50 2.5 45 2.25 40 ..I 2 _____ 35 D.-++ +::::L, 1. 75 ,.. 30 0 .p. 0 I _TI .__ L ,...1...-..L, 1.5 25 ____.1 I ·- L.l. L ..l 1.25 20 I ______!: ___ ;::.:!.!~! ______·r::-r· rl::i.. Lrl _!:~::;::~:!_____ ,..t=:L, . tt LJ I ------·----- _ 15 I B E F H N T 10 Lrl 5 ------B E F H N T Figure 5. Box plots of selected characters of Carex brevior (B), £. tenera var. echinodes (E), £. molesta (M), £. normalis (N), and£. tenera (T). Means indicated by heavy line; mode by capped line; and box indicates interquartile (25th to 75th percentile). -...... ) 0 Character 14 Inflorescence length 60 10 Ch""'"1: sp;k~".OO.L 55 0 I 0 9

50 0 I •-----· I 8 45 ·-----· 1 1 o 1 1 ·-----· 0 I I ~It.._ __t 40 ·-----· 6 I ll rr 11-•---··'-11~I I··-·-·· •-----·'tT L..l ·-----· Itt :: .IJ . ~~~r i .~~~ ., ~~~· : T 1 Jrt 0 Lr

~~ I i::i::iI 2 I I B E F H N T 10 ------·------·------·- B E F H N T gement 4 sp;ke "''", • , 3.75 Character 16 Inflorescence width/length ... 3.5 Chmot"II 0.9 3.25 0.8 3 . II.. .. 1 · 1 0.7 .J. 2.75 I.L...J...I 0.6 L.... J 2.5 l ...1·I I 2.25 0. 5 f I I 0 0.4 I.LLI . I . 1-E}I ._J_. 2 0.3 L. ..l Ll ...... I I.L._.LI I. 75 •--·--· 0.2 111~L .. ·-T· 1.5 1 l o.l ·-·------6------··1----·-····-F------f.1------·-N------·-r·-·------I. 25 . _____!:.::;: ______B E F H N T

Figure 5. Box plots of selected characters of Carex brevior (B),~.tenera var. echinodes (E), ~.molesta {M), £. normalis {N), and~.tenera {T). Means indicated by heavy line; mode by capped line; and box indicates interquartile {25th to 75th percentile). (table continues) --.....l Character 21 Spike width/length Character 23 Spike apex 1.2 8 6 1.1 5.5 5 0.9 4.5 I . . I .. 4 0.8 t. •-----· I 1-•-----·- I I ·-----· 0.7 I I r:bb I I 3.5 ~rr·-----·i--~--L~ I 3 0.6 ~ L_J . ·-----· H ·-----··-----· 1-·-----·-1 H ..6:..;..;..:.:1 ----- I . I 1____ _1 tt ·-----· 0.5 I ~Itt 2.5 I '-rl r-r::::r I ·r 2 0.4 • I Itt±± 8 ttl 0.3 1.5 B E F H N T 1 B E F H N T

Character 22 Spike shape 6 Character 24 Terminal spike: base shape 5.5 5 _r----r 5 -u······ ~ 4.5 4.5 4 Jl~H ·-----·I B 4 -----I 1-•-----•-1 "tt1-•-----•-1 I 3.5 L__l I 3.5 3 c:=:o·· .~.T . .. +± 3 2.5 r 2.5 2 nl 2 1.5 1.5 B E F M N T B E F M N T

~--

Figure 5. Box plots of selected characters of Carex brevior (B), ~. tenera var. echinodes {E), ~.molesta (M), £. normalis (N), and£. tenera (T). Means indicated by heavy line; mode by capped line; and box indicates interquartile {25th to 75th percentile). {table continues) -.) N Character 25 Lateral spike: base shape Character 28 Lateral spike: # male cycles 18 5 I ·-----· 4.5 16 4 14 3.5 12 .L 3 10 u~++I '······· 0 0 2.5 8 u 6 't I I .... . ~u·--'--•I I L...ll : I ...L. ll 4 L.. IJ.:..:..:.:.j_l I 1....1..-..1 ,J:::::L,;-:t=:t 2 1.5 ------~------·------·--·------tf___LJ ______I LTI LTI LTI I B E F H N T 0 B E F H N T

Character 26 Terminal spike: #male cycles 18 Character 30 Terminal spike: male cycles/female cycles 16 1.6 14 I 1.4 12 ·-----· 1.2 10 I J ___J, I 8 ·----· 1 0 :r I I 0.8 6 Jl__l_ I , __!__, Itt•-----• I IJ.:..:..:.:.j_l _ ~ o.61 tf.J.. ;~; I I I , llJ. 4 I·------1 L .. l 1-L--.,L 1 L.J ~ 0.4 ...... I 1 I I I· ••.••• 2 I l L .. l ----- I ----- r-:----t-i ----- I 0.2 I ·.::j::.· I IL:.LI 1-i:::::i.l L( 0 ·············------·------~------'------'------·------·--·· B E F H N T 0 ------·------·------______:::J::: ..... :::J::: ______,______B E F H N T

Figure 5. Box plots of selected characters of Carex brevior (8), ~.tenera var. echinodes (E),~-molesta (M), ~-normalis (N), and~-tenera (T). Means indicated by heavy line; mode by capped line; and box indicates interquartile (25th to 75th percentile). (table continues) -.Jw Character 32 Scale length (mm) Character 35 Scale width/Perigynia width 4.5 1.2 4.25 1.1 4 1 3.75 0.9 3.5 .L 0.8 ___L I .J.. .J.. 3.25 p __L. 0.7 3 0.6 +:+~-LJ-1 +=++4 ..l. .J .. 2.75 . I +:+._J_i ~4~ 0.5 t...L...... l...l I I I...L...... l...l I --( r' 1~1 1-:::::::_1 1-'------1 I t_____ t 2.5 ·-T L=r Itt Lrl 0.4 I 2.25 0.3 I ·r 0 2 B E F H N T 1. 75 B E F H N T

5.5 Character 36 Perigynia length (mm) 5 _ t Character 34 Scale length/Perigynia length 1 3 4.5 ___L. 4 r 1-:::l--· 1.2l 8 IL:.:l.. L:T 1-i:::::i-1 3.5 L_ ___ll 1.1 0 _ __I___ I I.LLI l I 3 ..l. t__ ___l ..l. 1?1 I 1------1L_ __j 0.9 8 2.5 I J::::tL . .l IL:LI I I I ltj-1 0 8 2 I · I ---~--l.t.:.:.:.:LI 11:~~:11 I y I 0. 7 _L._ 1._ _.1 T:":T B E F H N T 0.6 0 I LTII 1 I

0. 5 ------·------·------·------·------~------· B E F H N T

Figure 5. Box plots of selected characters of Carex brevior (B), ~. tenera var. echinodes (E), ~. molesta (M), ~. normalis (N), and~.tenera (T). Means indicated by heavy line; mode by capped line; and box indicates interquartile {25th to 75th percentile). (table continues) ~ Character 37 Perigynia width (mm) Perigyn~as~:pe.. width/1 ength 3 .4 Ch.,,,,., 40 ,, .. '''· I

3.~ I 8 I 1._ ·-----· ·----- I 7.5 1-'-----'-1 ++1-·-----l H ~::J:::t I 2.4 tr r---i 2 ,; IT 'L ·~ I I tr I l ··n·¥ 1.8 8 •-----·I 1-'---··'·1•--·-·· I ·-----· I 0 IL-J..I ++ •-----· ~ 1...... _.._1 ~:: : Lrl ~..( 1-·=r··l ~~_.~~ I ·--~--

1.2 --·------·------·------·------·------1 B E F M N T

·------·------·------~- 8 E F M N T

Character 41 Perigynia length to widest 0.5 point/perigynia length Character 39 Perigynia width/Perigynia length 0.475 0.9 0.45 0.425 0.8 ...L. 0.7 l..l..-..LI 0.4 1... . .1 0 .J. 0.6 181 .__!__, l 0.375 ----- t ~----~ I 1-1·---·-· IL-f-1 i ..J.. I 0.5 T I L . .l ... ..• ··-·--· 0.35 M=+: I I I I 0.4 ! 1--T--1 ··-r 0.325 I.LJ_Ig I n 0.3 LT ..L. I.TI L __l LJ. I I II 0.3 ·------·------·------·------·------· B E F M N T 0.275 lnl 0 I I L.J I ---,-- 0.25 0.225 I.TI 0 I 0.2 B E F M N T

Figure 5. Box plots of selected characters of Carex brevior (B), ~. tenera var. echinodes (E),~.molesta (M), ~. normalis (N), and~.tenera (T). Means indicated by heavy line; mode by capped line; and box indicates interquartile (25th to 75th percentile). (table continues) U! Character 43 Beak length (mm) Character 50 Achene length (mm) 3 0 2 2.8 0 1.9 2.6 1.8 J. I I 0 ,L_L...... 0 •-----· 2.4 .J.. I 1.7 ,l__l_, L ___] ,l__l_, I ,j___J_, 2.2 L_.l ,g, 1.6 I I ·-----·-1 ]_____] 2 L. j I I I 1.5 1 1.8 .L I ,r:::c r+d, J:::J. I 1.4 T I J:::t 1.6 I 1.3 L ..l ...L. 0 TT,L~L 1.4 ,J..__J_, 1.2 1... .. 1 I ~,~ ~T 1.2 I 1.1 I B E F H N T 0.8 +··------•------· B E F H N T

Character 51 Achene width (mm) 1.7 Character 44 Beak length/Perigynia length 1.6 0.8 1.5 ,1._.Ji ____, 0.75 1.4 0.7 1.3 0.65 .J.. 1.2 T ,.L .. l., I 0.6 1.1 ...... Lr ...... ··-'·-· 0.55 ,_:::L_, ,J::L_, ...L_. .J. 1 r~r ~ I ~ ,.!..... !., 0.5 I ""L":T I ""L":T r+=fr rf=tr 0.9 . I I I 0.45 r~rI r-F::j-r I I I 0.8 0.4 B E F H N T 0.35 ------~~~(...... J...... ~---·····---·--·-···-·--•---·····-·· 8 E F H N T

Figure 5. Box plots of selected characters of Carex brevior (B), £. tenera var. echinodes (E), £. molesta (M), £. normalis (N), and£. tenera (T). Means indicated by heavy line; mode by capped line; and box indicates interquartile (25th to 75th percentile). (table continues) 0\-...... l Character 52 Achene width/Achene length Character 55 Achene length/Perigynia length 0.7 0.95 0.65 0.9 0.6 0.85 Ji 0.55 ...L. I L.... ·I I I 0.8 : I I 0.5 +:~+ ~-[}~ 0. 75 I I •-----·•-----· 0.45 I l ...L. IQ 1.__._1...L. ·-----· 1...1..-....Lt I 0.7 I ·-----·ILl. ,..!._._!,ILl_ 0.4 I ·-T-· ITI··r· ~ '·Loo!·'L. . .J ' ' C l' J.... .J ' 8 0.65 I 0.35 0.6 0.3 0.55 ''r'' I I T I B E F H N T 0.5 B E F H N T Character 56 Achene width/Perigynia width 0.95 0.9 Character 53 Achene shape: width/length (see Fig. 1) 0.85 8 0.8 7.5 0.75 7 0.7 1 • I 6.5 0.65 ----I _o----I ·-----·I 6 1 1 I.,,.J.,tt,.J.,r1 0.6 l..l.--LI·-----· 0 1-"---- L:LI I I I I 5.5 ±=ttt ++ ~ 0.55 I L I I 8-.....! L.l. .l l l II H I ~.LJ.----- 5 I 1-•-----•-t ------1---, 0.5 r I 4.5 0.45 . ···rI 1 4 0.4 3.5 B E F H N T 3 B E F H N T

Figure 5. Box plots of selected characters of Carex brevior {B), ~-tenera var. echinodes {E), ~-molesta {M}, ~-normalis {N}, and~-tenera {T}. Means indicated by heavy line; mode by capped line; and box indicates interquartile {25th to 75th percentile}. {table continues} -.....) -.....) 78 because of their large sample sizes. The remaining taxa have much smaller sample sizes (Table 5), nonetheless, their character distributions tended toward normal bell-shaped curves. High values for the Shapiro-Wilk statistic, a measure of frequency distribution shape, indicate that characters of taxa with smaller sample sizes were normally distributed; however, these estimates were not always significant (I! < 0.05). The lack of significance inplies that while the dataset tends toward a normal distribution, it may not provide a true estimate of the mean or include the entire range of variation for some characters of C. normalis,

C. festucacea, C. tenera, and C. tenera var. echinodes. Many of the qualitative characters also had normal distributions because character states were ordered along a gradual scale where intermediate values could be used. Based on these results, it was decided that multivariate analysis could be performed by parametric methods.

Infraspecific variation. Most of the frequency distributions for continuous characters were normal bell-shaped curves indicating that no further infraspecific subgroupings were necessary within the _c. brevior group. Also, normal distributions for the entire dataset of each taxon indicates there are no significant differences between populations. This is compatible with the results obtained from multivariate analysis.

Univariate tests of taxonomic si~~:nificance. Pairwise comparisons of taxa using the the students t-test was not useful in selecting distinguishing characters for this dataset. For example, _c. molesta and ..C. brevior have significantly different means for 52 of the 56 characters analyzed (I! < 0.0001). However, 79 frequency distributions for all the characters overlapped extensively (Table 6 and

Figure 5). Consequently, no single character can reliably separate this difficult pair of taxa. Likewise, .C. festucacea (N = 30) and .C. tenera (N = 58) are also a troublesome pair to distinguish with only 24 characters having significantly different means (12 < 0.0001) yet with overlapping ranges of variation. The t-test comparisons of other taxa yielded similar results: separate character means but overlapping variation. The extensive overlap of taxonomic characters made it difficult to separate taxa on the basis of univariate statistics alone. Multivariate analysis was necessary to identify character combinations capable of maximizing differences between taxa.

Multivariate Analyses

Canonical Discriminant Analysis (CFA) plots specimens according to a classification variable using canonical values in a manner similar to Principal

Component Analysis. The discriminating power of a canonical axis is a proportion of the total variation of the entire dataset explained by variables correlated to that axis. Specimens are assigned a canonical value for each canonical axis. A canonical value of a specimen represents the combined

character states for suites of variables correlated along a particular axis.

Specimens can be plotted in n-dimensional space depending on the number of

canonical axes.

Multivariate tests of taxonomic si~nificance. The SAS CANDISC

procedure plotted the randomly collected specimens of the _c. brevior group into

six distinct taxon clusters along four canonical axes (Figures 6-8). The Wilks' 80

Lambda Test determined that the six clusters are significantly different with an F­ value of 24.08 (p < 0.0001). Multivariate analysis suggests that .C. brevior, .C. festucacea, .C. molesta, _c. normalis, _c. tenera, and .C. tenera var. echinodes are distinct morphological forms and justifies their recognition as separate taxa.

Character variables which are highly correlated may actually be measuring the same features of the plant. This would essentially double-weight the discriminating power of that particular morphological feature. Upon examination of DISCRIM Within-class Correlation Coefficients, I found 11 character pairs which were highly correlated with the absolute value of r 2.. 0.7500. One character of each pair was removed and the dataset was reanalyzed by discrimant and canonical methods. The taxon classes were still significantly different with a slightly higher F-statistic of 26.75 (P < 0.0001), indicating that the removed characters were more variable within the taxa than between them.

While measuring vegetative characters 9-12 in the laboratory, I determined these features have limited taxonomic usefulness for the .C. brevior group. This was confirmed by DFA when low F-values for vegetative characters 9-12 were obtained, thus indicating these characters had little discriminating power for separating taxa. In an effort to save time, I began evalutating vegetative culms only on every 5th specimen. However, multivariate analysis rejects samples with missing variables, and if characters 9-12 were included in multivariate analysis, then my total sample size would have been reduced to one-fifth of the original sample size. Therefore, characters 9-12 were removed from the final multivariate analysis in order to increase sample sizes. 81

Infraspecific variation. To determine if there were any morphological differences between populations within a taxon, the same canonical plots were re­ illustrated to identify specimens by their population codes (Table 5). The distribution of population codes within taxon clusters is important. Specimens of each population were found throughout taxon clusters in all three canonical plots

(Figures 9-11). No significant patterns within any of the clusters were observed, indicating that no further subgroupings are necessary within species of the _c. brevior group. The distribution of population codes also indicates that most of the total morphological variation expressed by Iowa taxa occurs within populations.

Specimens of sympatric populations occupy canonical positions in between taxon clusters and are predominately found in hemispheres of the taxon clusters closest to the sympatric species (Figure 12). Removal of specimens collected at sympatric sites reduces morphological variation, eliminates overlap of taxon clusters, and results in higher F-values of significance. This is readily observed in sympatric populations of _c. brevior and _c. molesta. It is evident that some portion of the total morphological variation within Iowa taxa is found only among sympatric populations.

Taxonomic Characters

Suites of morphological characters are correlated to each canonical axis which are orthogonal to one another. All the canonical axes separate _c. brevior and _c. molesta from the rest of the taxa. Morphological characters correlated to

Canonical axes 1 and 2 distinguish .,C. brevior from Carex molesta. Canonical axis M 6 M

M M HMMM MMM MMMMMM M 4 + M M MMMMMM M MMM MMM M M MMMMMMM MMM M M CAN2I M M MMM MMM M M M M M MM MM M MMMMMMMMMM M M M 2 + BMM M MM M N 8 M M M N H NN N H N M 8 F N N 8 M 8 M M F H 8 8 N EN H 8 8 H NFN NN NN 0 8 H EN E + 8 8 F F F E E EN 8 88 8 8 F F 8 888 88 F F NF 8888888888 8 F FF 8 888888 8 F F FF F FF T EE b 88 8888 8 8 T F F ET E T -2 + 8 8 888888888 T E 8 8 888 8888 8 88 F T F T 8 888 888888 8 8 T T TT T T 888 8 8 8 T TTTTTT 8 88 88 8 T TT T T 8 8 88 8 8 T T T TT 8 T T T T T 8 T -4 + T T

8 T T T

-6 ---+------+------+------+------+------+------+------+------+--T -8 -6 -4 -2 0 2 4 6 8 CANI Figure 6. Plot of canonical axes 1 and 2 showing ordination of Carex brevior (B), Carex festucacea Carex (M), Carex Carex and Carex 00 (F)' molesta normalis (N), tenera (T), tenera var. echinodes N (E). Taxon clusters were significantly different at F = 24.08. 8 EE E E E E 6 EE E E E CAN3 E E 4 E H 8 T E H 8 888 8 H M 2 H M M 8 8 8 H M B T 8 88 B B H HH H M M""' M B 8T 88 88 B 8 BB B H H ""' M M M T B BT88 BBBBB 8 B HH B B MMMM M M M B TBBT B BB BBBBBB B H B ""'""'"""""M M M 0 BB BBB BBBBBBBT BB BH B H B HNMM~M""'M BTTBT BB BB BB BH B HH H B N M B M M M MMMM M M M T B 88 BB TTTB 8 H H B MM """""""'MM T TB TT T B B M M M M M"""""""'M M T T T T B BB B H N M ""'M MMMH T FB M -2 T F B ""'M T T T F F F M T M F F F F F T F F -4 F FF F F F FF F F -6 F F

-8 ---+------+------+------+------+------+--. -6 -4 -2 0 2 4

CAN2

Figure 7. Plot of canonical axes 2 and 3 showing ordination of Carex brevior (B), Carex festucacea (F), Carex molesta (M), Carex normalis (N), Carex tenera (T), and Carex tenera var. echinodes 00 (E) • Taxon clusters were significantly different at F = 24.08. (.;)

---'~ 6 N N N N N N N N N N N N NN 4 NN N N N N N CAN4I F N F FF F N F F F N 8N B 8 F 8 8 88 N 2 + F 8 8 8 8 F FFF F F N888B 8 8 F F B 8118 8118 F 8 88 B 88888 E F F F M M 888888 M 8 M M 8 88B8MB8 8118M E F F M B BB 8BBBM8BBBB BE 8 EE 0 F M M MMM88BBB8118BBMB 8 8T MMBM8118118MBBM 888 E E E M 88MM8B8118 M B E E B M MM BBMB811B 8 E MMMT 8M MB MB MMTMM 88 TM M TM TMMBM MMM -2 TMT MMTTIIM811T8 M T TT TMMM M T TTTT MT T 8 T M M MT T TT M T T T -4 T T T

-6 ---+------+------+------+------+------+------+------+------+------+-- -8 -6 -4 -2 0 2 4 6 8 10 CAN3

Figure 8. Plot of canonical axes 3 and 4 showing ordination of Carex brevior (B), Carex festucacea (F), Carex molesta (M), Carex normalis (N), Carex tenera (T), and Carex tenera var. echinodes 00 (E). Taxon clusters were significantly different at F = 24.08. ~ a 6 m

m m n hn n nba 1111111mam m 4 hn aammam nnh nhm a b nnmh jnb ama b m CAN2 a aaabbmh b ann nnm ha b nhhohnnmmb b II Ill 2 bnan mn a Ill g 111a II c c cc p c f h h a c c d 0 b h g 0 c 0 g c ec k h g c cap cc cc e 0 + h c ek e g 0 a a a e e em e 1 1d 0 g a a k j hdo 1g a a ca e 11o g1 1 g o od a aa e d 11 im o d g a a a a a aa kee g 11 o111 g 1 p a a ek e k -2 + 1 0 1 o1 g 11 g o 1 k e 1 1 11o hoo1 1 1d a k a k 1 1 1 1 1d goo o 0 0 k p pk k k oo1 0 1 0 p k p kp p k og 00 d p pp p k 1 1 o1 b k p p pp 1 k k p p p -4 ! 0 p p p

p p p

-6 ---+------+------+------+------+------+------+------+------+-- -8 -6 -4 -2 0 2 4 6 8 CANI

Figure 9. Plot of canonical axes 1 and 2 showing ordination of specimens according to population codes of collection sites (Table 5) (Compare with Figure 6). 00 Vl 8 •• • • • • 6 •• • CAN3 •• • 4 • • p p c o1h 1 • c n 2 c m B g 1 o c m p o 11 o 1 c cc m b m Hn m o ok oo 11 1 1 g1 g p c mm b b h p 1 op11 llglg o o cc d g oan b b h m o plop o 11 g11111 g c b nm nh mamb j a H 0 11 dog oohdll ok og dm h c g cf n m hnnhb m nmn bpkip od o1 go he o kc c o c a b m n n nnn n H n m p p o oo 11 kpkd 1 c c g n a hnaa ah m n p pg kk k o 1 h m a a a hnnn mm a a p p p k g gd o k c b hn a h aag k a1 H -2 k a 1 aa a p p p a a 0 • a • • • a -4 p a • a a • a a a a a a • -6 • • -8 ---+------+------+------+------+------+-----. -6 -4 -2 0 2 4 CAN2

Figure 10. Plot of canonical axes 2 and 3 showing ordination of specimens according to population codes of collection sites (Table 5) (Compare with Figure 7). 0\00 6 t c c c c c c c c c c c c c c cc 4 cc + c m k m p CAN4I a c a aa a f a a a k 1c m d a g h g1 p 2 + a 0 g I o a aaa a a cbgdd g 1 a a 0 hn g 1a1 a goo o o11d1 e a a a h m b o1 gd I a I n a o 1ob1go1 !Jill n e e a a a d t 1 ohoomgo1ot le 1 ee 0 o a b mhago odllmg1omo 0 hp e + gao nobonobilb 11 t E e e h i tnmo1 1bt h I e E e I a ma o1m11o 1 1 E andc. gn mo bt e mapm m ot km m ka knhta mbm -2 + kak hmpkhn hako m p kk pnom a P kpkp mp p 0 p h h np k pk m p k p p p -4 p p p

-6 ~---+------+------+------+------+------+------+------+------+------+-- -8 -6 -4 -2 0 2 4 6 8 10 CAN3 Figure 11. Plot of canonical axes 3 and 4 showing ordination of specimens according to population 00 codes of collection sites (Table 5) (Compare with Figure 8). -..1 6 .. a

a a .... "' •. 4 ...... aaa a ..a e ...... a .. a .. e ••9 • CAN2 a ..... a.. .. e ...... •9• ..

2 .. ..

• .. e .. • ...... • . ~ .. 0 ...... e ee • .. .. e ..a ...... ea .. •e e • e ...... a ee ee .. e .. .., ...... ee ..aae • .. .. -2 e • e •9 .. ee .. .. a e ea ...... a a ae .. e e e e ee ••• • • • • ••8 .. a ...... e 9 9 •9 • 9 a -4 .. a

-6 ---+------+------+------+------t------t------t------t------T-- -8 -6 -4 -2 0 2 4 6 8 CANI

Figure 12. Plot of canonical axes 1 and 2 ordinating specimens of Carex brevior, Carex molesta andCarex festucacea by whether they were collected at allopatric sites (e) or sympatric sites (Compare 00 (•) (Table 5). Lines delimit boundaries of intermediate specimens with Figure 6). 00 89

3 separates _c. festucacea from _c. tenera var. echinodes; and canonical axis 4 separates _c. normalis from _c. tenera.

The SAS CANDISC procedure provides total-sample correlations between the canonical coefficients and the character variables. Canonical characters with the highest absolute value are the most significant characters for that particular canonical axis. Eigenvalues are measures of significance for canonical axes. They represent the ratio of between-class variation to the within-class variation of the morphological characters corresponding to a particular cannonical axis.

Eigenvalues for each canonical axis indicate the proportion of the total variation attributed to those characters that are correlated to a particular axis (Table 7).

Table 7. Eigenvalue and the percentage of the total variation explained by each canonical axis. Canonical Axis Eigenvalue % Total Variation Can1 10.57 45% Can2 6.03 26% Can3 2.82 12% Can4 2.66 11%

Stepwise Discriminant Analysis. The Stepwise Discriminate Analysis selects taxonomic characters on the basis of their overall discriminating power as measured by the F-statistic. Morphological characters are correlated to Euclidean vectors between taxon clusters plotted in n-dimensional space. Morphological variation of a character is represented by vector length. Significance of each character is determined by the ratio of the average variation between taxa to the average variation within taxa (see Table 8). 90

Table 8. Stepwise discriminant characters separating Carex brevior, ..C. festucacea,. ..C. molesta, .C. normalis, .C. tenera and _c. tenera var. echinodes.

Character Label Partial R2 F-Statistic1 51 Achene width 0. 7170 183.386 16 Inflorescence width/length 0.6591 139.564 8 F. culm upper leaf width 0.4834 67.380 36 Perigynia length 0.4357 55.431 29 Lateral spike female cycles 0.3938 46.523 7 F. culm upper leaf length 0.3015 30.817 48 Dorsal nerve number 0.2441 22.996 14 Inflorescence length 0.2333 21.610 17 Spike number 0.2590 24.748 24 Terminal spike base shape 0.1721 14.672 28 Lateral spike male cycles 0.1460 12.034 32 Scale length 0.1378 11.222 55 Achene 1./perigynia length 0.1350 10.923 45 Beak shape 0.1136 8.944 4 Fertile culm height 0.1099 8.597 38 Perigynia 1. to widest point 0.0980 7.536 41 Perigynia 1. tow. pt./1. 0.1260 9.976 47 Ventral nerve appearance 0.0871 6.585 46 Ventral nerve number 0.0829 6.223 42 Perigynia shape 0.0797 5.939 2 Fertile culm width 0.0839 6.262 35 Scale w.jperigynia width 0.0683 4.998 43 Beak length 0.0664 4.836 33 Scale width 0.0614 4.436 39 Perigynia width/length 0.0629 4.535 37 Perigynia width 0.1312 10.182 34 Scale 1./perigynia length 0.1298 10.024 52 Achene width/length 0.0745 5.395 22 Spike shape 0.0405 2.822 26 Terminal spike male cycles 0.0398 2.761 15 Inflorescence width 0.0399 2.759 40 Perigynia shape (l.jw.) 0.0429 2.970 27 Terminal spike female cycles 0.0365 2.502 20 Spike width 0.0356 2.429 44 Beak 1./perigynia length 0.0288 1.947 19 Spike length 0.0272 1.834 23 Spike apex shape 0.0260 1. 749 1Characters are arranged in descending order of significance by Wilk's Lambda (not given). The F-statistic is provided for comparison. 91

Table 8 lists morphlogical characters selected by SAS STEPDISC procedure for separating all six taxa of the ..C. brevior group. Characters are ranked in descending order of significance according to Wilk's Lambda. The

F-statistic, another measure of significance, is provided because it is more commonly used. The partial R2 is a proportion of the morphological variation explained by the character. Unknown specimens can be identified by sequentially comparing discriminate features of the plant with the box plots and minimum/maximum values of the dataset (Figure 5 and Table 6). For construction of dichotomous keys, STEPDISC identified key characters for segregating subgroups within the .C. brevior group. Canonical plots show that C. brevior and C. molesta can be grouped together for comparison to the rest of the group. Table 9 lists characters which separate ..C. brevior and .C. molesta from .C. fesrucacea, .C. normalis, .C. tenera, and ..C. tenera var. echinodes. Table 10 lists characters separating ..C. fesrucacea, ..C. tenera var. echinodes, _c. ten era and _c. normalis into two pairs. STEPDISC also provided character suites to separate difficult pairs of taxa (Tables 11-14).

Morphological Descriptions

Discriminant characters provided by STEPDISC were combined into character suites to create species descriptions. These character combinations will separate taxa within the ..C. brevior group when considered together. These descriptions include means and interquartile ranges (compare with Figure 5 and

Table 6). 92

Table 9. Stepwise discriminant characters separating the .C. brevior group into two subgroups for dichotimous key construction: Subgroup A = .C. brevior and .C. molesta; Subgroup B = .C. festucacea, .C. normalis, .C. tenera and .C. tenera var. echinodes. Character Label Partial R2 F-Statistic1 37 Perigynia width 0.6040 558.312 14 Inflorescence length 0.3039 159.329 48 Dorsal nerve number 0.1700 74.547 17 Spike number 0.1080 43.963 16 Infloresence width/length 0.1185 48.599 47 Ventral nerve number 0.0505 19.199 24 Terminal spike base shape 0.0535 20.339 32 Scale length 0.0524 19.841 50 Achene length 0.0331 12.272 35 Scale widthjperigynia width 0.0321 11.845 51 Achene width 0.0295 10.818 23 Spike apex shape 0.0025 9.967 20 Spike width 0.0272 8.938 7 Fertile culm upper leaf length 0.0246 12.168 29 Lateral spike female cycles 0.0242 8.758 41 Perigynia 1. tow. pt./length 0.0199 7.138 44 Beak length/perigynia length 0.0247 8.882 33 Scale width 0.0146 5.200 56 Achene widthjperigynia width 0.0179 6.355

1Characters are arranged in descending order of significance by Wilk's Lambda (not given). The F-statistic is provided for comparison.

~ molesta: 3-5 globose spikes with spreading beaks, spike arrangement closely aggregate to spreading at base of inflorescence, rachis stiffly erect; lateral spike base shape round with few male florets (2-3 male cycles on lateral spikes); perigynia lanceolate to ellipitic; 3-4 ventral nerves; scale length/perigynium length

0.69-(0.71)-0.79, perigynium 3.08-(4.06)-4.25 mm long, 2.17-(2.32)-2.50 mm wide; beak 2.00-(2.16)-2.33 mm long, beak length/periginium length 0.50-(0.53)-0.56, beak shape tapered; achene 1.08-(1.16)-1.25 mm wide, achene shape ovate; leaf width< 4 mm. 93

Table 10. Stepwise discriminant characters separating Carex festucacea, _c. normalis, .,C. tenera and .,C. tenera var. echinodes into two pairs: Pair 1 = .C. normalis and .C. tenera var. echinodes; Pair 2 = .C. festucacea and .C. tenera. Character Label 1 36 Perigynia length 2 Fertile culm width 38 Perigynia length to widest point 8 Upper leaf width 23 Lateral spike female cycles 31 Lateral spike male/female cycles 23 Spike apex shape 32 Scale length 4 Fertile culm height 46 Ventral nerve number 41 Perigynia length to widest pointjperigynia length 15 Inflorescence width 47 Ventral nerve appearance 45 Beak shape 39 Perigynia width/length 35 Scale width/perigynia width 4 Spike number 1Characters are arranged in descending order of significance by Wilk's Lambda (not given).

Carex brevior: 3-5 spikes varying from globose to obovate with ascending beaks, spike arrangement loosely aggregate to strongly moniliform, rachis erect to

nodding; lateral spike base shape tapered to strongly clavate with several male florets (3-6 male cycles on lateral spike); perigynia subrotund to circular, no ventral nerves or nerved at base only; scale length/perigynium length 0.86-(0.92)-

0.98; perigynium 3.41-(3.67)-3.9 mm long, 2.42-(2.56)-2.75 mm wide; beak 1.50-

(1.71)-1.83 mm long, beak length/perigynia length 0.42-(0.46)-0.49, beak shape abruptly constricted; achene 1.33-(1.41)-1.5 mm wide, achene shape oval to ovate; leaf width < 4 mm. 94

Table 11. Stepwise discriminant characters for separating Carex brevior and _c. m~ta. Character Label 1 45 Beak shape 24 Terminal spike base shape 34 Scale lengthjperigynia length 51 Achene width 29 Lateral spike female cycles 7 Fertile culm upper leaf length 46 Ventral nerve number 16 Inflorescence width/length 43 Beak length 35 Scale widthjperigynia width 26 Terminal spike male cycles 15 Inflorescence width 20 Spike width

1Characters are arranged in descending order of significance by Wilk's Lambda (not given).

Table 12. Stepwise discriminant characters for separating Carex festucacea and ..C. tenera. Character Label 1 42 Perigynia shape (position of widest point) 22 Spike shape 26 Terminal spike male cycles 2 Fertile culm width 50 Achene length 8 Fertile culm upper leaf width 15 Inflorescence width 27 Terminal spike female cycles 47 Ventral nerve appearance 48 Dorsal nerve number 38 Perigynia length to widest point (mm) 51 Achene width 52 Achene width/length 4 Fertile culm height 17 Spike number 32 Scale length 33 Scale width 39 Perigynia width/length 45 Beak shape 95

Table 13. Stepwise disciminant characters for separating Carex brevior and _c. festucacea. Character Label 1 29 Lateral spike female cycles 51 Achene width 26 Terminal spike male cycles 47 Ventral nerve appearance 33 Scale width 17 Spike number 38 Perigynia length to widest point 14 Inflorescence length 32 Scale length 21 Spike width/length 27 Terminal spike female cycles 48 Dorsal nerve number 19 Spike length

1Characters are arranged in descending order of significance by Wilk's Lambda (not given).

Table 14. Stepwise discriminate characters for separating Carex normalis and C. tenera var. echinodes. Character Label 1 8 Fertile culm upper leaf length 40 Perigynia shape (width/length ratio) 48 Dorsal nerve number 32 Scale length 4 Fertile culm height 2 Fertile culm width 20 Spike width 1 Fertile culm leaf number 27 Terminal spike female cycles 3 Fertile culm leaf extended length 38 Perigynia length to widest point 18 Spike arrangement 1Characters are arranged in descending order of significance by Wilk's Lambda (not given). 96

Carex normalis: 4-10 spikes varying from ovoid to obovoid with spreading beaks, spike arrangement aggregate to spreading at base, rachis stiffly erect; lateral spike base usually round to tapering with few male florets (2-4 male cycles on lateral spikes); perigynia narrowly ovate, 3-5 ventral nerves; scale length/perigynium length 0.71-(0.75)-0.79; perigynium 3.08-(3.42)-3.71 mm long,

1.58-(1.75)-1.92 mm wide; beaks 1.50-(1.72)-1.92 mm long, beak length/perigynia length 0.47-(0.50)-0.53, beak shape tapered; achene 1.0-(1.04)-1.08 mm wide, achene shape elliptic to ovate; leaf width > 4 mm.

Carex festucacea: 4-9 spikes varying from subrotund to obovate with

spreading beaks, spike arrangement moniliform, rachis nodding, lateral spike base

clavate with numerous male florets (5-12 male cycles on lateral spikes); perigynia

round to elliptic, 3-4 ventral nerves; scale length/perigynium length 0.82-(0.88)-

0.94; perigynium 2.75-(3.04)-3.17 mm long, 1.67-(1.80)-1.92 mm wide; beak 1.25-

(1.35)-1.50 mm long, beak length/perigynium length 0.42-(0.44)-0.47, beak shape

abruptly constricted; achene 1.0-(1.05)-1.08 mm wide, achene shape elliptic; leaf

width< 4 mm.

Carex tenera: 3-7 spikes varying from ovoid to oblong with ascending

beaks, spike arrangement varying from aggregate at inflorescence apex to

moniliform, rachis weak nodding, flexuous, often zig-zagging; lateral spike base

tapered with few male florets (3-6 male cycles on lateral spikes); perigynia

broadly ovate, 4-5 ventral nerves, scale length/perigynium length 0.67-(0.94)-1.0;

perigynium 2.75-(3.02)-3.12 mm long, 1.50-(1.65)-1.75 mm wide, beak 1.33-(1.52)-

1.67 mm long, beak length/perigynium length 0.47-(0.5)-0.53, beak shape tapered;

achene 1.0-(1.03)-1.08 mm wide, achene shape ovate; leaf width -=5. 3 mm. 97

~ tenera var. echinodes: 5-8 spikes varying from ovoid to oblong with widely spreading beaks (echinate), spike arrangement strongly moniliform, rachis

nodding, flexuous, or often zig-zagging, lateral spike base tapered with very few

male florets (2-4 male cycles on lateral spikes); perigynia narrowly lanceolate to

ovate, 3-4 ventral nerves; scale length/perigynium length 0.62-(0.66)-0.69;

perigynium 3.91-(4.18)-4.42 mm long, 1.67-(1.74)-1.83 mm wide, beak 2.08-(2.27)-

2.42 mm long, beak length/perigynium length 0.51-(0.54)-0.56, beak shape

tapered; achene 1.01 mm wide, achene shape narrowly ovate; leaf width ~ 3 mm.

Intermediate Specimens with Mixed Character Suites

Typically, suites of certain character states are highly correlated to a

specific taxon, e.g., compare .C. molesta with .C. brevior. However, at sympatric

sites, specimens growing between different taxa tend to have mixed character

suites, i.e., they display mixed character states not usually found together on one

specimen.

Collection numbers of specimens with mixed character suites are presented

in Table 15. These specimens are found in canonical plots in intermediate

positions between taxon clusters and are identified as "intermediate specimens".

Intermediate specimens were assigned mixed classification probabilities by the

SAS DISCRIM Posterior Probablility of Classification Error. I disagreed with

SAS DISCRIM classifications of some specimens, because I assigned different

weights to certain characters (these specimens are identified by a"**").

Specimens with primary classification probabilities less than 0.9999 were collected

at sympatric sites. I could detect mixed character suites in specimens with 98

Table 15. Specimens displaying mixed character suites as identified by (1) mixed classifications of SAS DISCRIM1 procedure; (2) intermediate location between taxon classes on canonical plots; and (3) observation of atypical character combinations. Taxon Site DISCRIM-Classifications4 Coordi nates5 Code 2 Code 3 Posterior Probabilities CAN1 CAN2 8:104 * BSM ! 8:0.9966 T:0.0034 -0.255 -2.526 8:110 BSM ! 8:0.9998 M:0.0002 -2.722 -0.170 B:120b BSM ! 8:0.9985 M:0.0015 -2.691 -0.338 8:194 CHSP 8:0.9999 M:0.0001 -1.446 -1.268 8:292 PKSP ! 8:0.9999 M:0.0001 Not Plotted 8:295 PKSP ! 8:0.9999 M:0.0001 -2.942 -2.340 8:319 * PKSP ! 8:0.9954 M:0.0046 -3.756 0.646 8:327 * PKSP ! 8:0.9554 M:0.0451 -2.414 0.419 8:318 ** PKSP ! M:0.9966 8:0.0044 -0.247 1.166 8:520 ** WP ! M:0.9029 8:0.0971 -1.277 0.977 8:530 ** WP ! M:0.9996 8:0.0004 -1.027 1.857

M:101b * BSM ! M:0.9494 8:0.0506 -1.914 0.763 M: 137 OTC ! M:0.9997 8:0.0003 -1.017 1.979 M: 147 OTC ! M:0.9998 8:0.0002 -0.868 2.074 M:160a M:0.9999 8:0.0001 -0.284 1.628 M:308 * PKSP ! M:0.9786 8:0.0214 -0.879 0. 724 M:314 * PKSP ! M:0.9804 8:0.0148 -1.557 1.065 M:328 * PKSP ! M:0.9852 8:0.0148 -0.366 0. 776 M:906 BSM ! M:0.9993 F:0.0007 2.441 1.906 F:101a * BSM ! F:0.7575 M:0.2425 1. 512 0.863 F:894 BSM ! F:0.9999 8:0.0001 1.768 1.283 8:945 ** BSM ! F:0.9998 8:0.0002 0.400 -1.316

T:055 * SRCP ! 6 T:0.9671 E:0.0329 2. 776 -2.678 1SAS DISCRIM Posterior Probability Error-rate Estimates. 2Letter represents Taxon code with collection number (Tab. 1). (*) marks specimens with DISCRIM classification of P 2.. 0.0034). (**) author classified specimen differently than SAS DISCRIM. 3See site identification codes (Tab. 2). (!) denotes sympatric site. 4First column represents DISCRIM posterior classification of the OTU by Taxon code followed by probability of certainty. Second column represents probability that the OTU should be classified into second taxon. 5Canonical coordinates for axes 1 and 2. 6Specimen collected near ..C. normalis and ..C. tenera. 99 secondary classification probabilities greater than 0.0034 (identified with a "*").

Specimens with mixed SAS DISCRIM probabilities less than 0.0034 tended to display character states at the extreme tails of univariate frequency distributions for that taxa. The habitats of intermediate specimens were greatly disturbed sites: sandy river raceways (PKSP); flooded prairie swales between dry knolls

(WP); eolian sand dunes (BSM); or exposed subsoil (BSM).

Intermediate Specimens of Carex brevior and Carex molesta

Specimens B:318, B:520 and B:530 ("B" =_c. brevior) occupied intermediate positions between taxon clusters of C. molesta and _c. brevior. They are atypical with mixed characters. Their primary SAS DISCRIM classification was _c. molesta, but other features of their perigynia and inflorescence are more representative of _c. brevior. DISCRIM classified B:318, B:520 and B:530 as C. molesta because they had shorter scale lengths; narrowly ovate achene shapes with shorter achene lengths; short spike lengths; high spike width/length ratio (i.e. round spikes); round terminal spike bases; few male florets on lateral spikes; spikes aggregated on inflorescence; low ratio: male cycles/female cycles on lateral spikes; low ratio: achene width/perigynia width; strongly to finely nerved ventrally; high dorsal nerve count; and tapering beaks. I classified them as C. brevior because of the subrotund shape of the perigynia and ascending beaks.

Many perigynia had no ventral nerves. These specimens were found in typical .C. brevior habitats on elevated knolls (B:520 and B:530) or well-drained sand

(B:318). 100

Mixed character suites for B:292 and B:295 include .C. molesta features with round, approximate spikes; ellipical perigynia, tapering beaks, strongly ventral nerves; and .C. brevior features with lateral spikes with tapering bases and high ratio: male cycles/female cycles; long achene length; high scale length to perigynia length ratio; and they are shorter plants with short leaves. To me they resemble .C. brevior more than .C. molesta. Mixed character suites for M:308 and M:314 ("M" =.C. molesta) include .C. brevior features with moniliform spikes with ascending beaks and moderatly clavate or tapering bases. However, perigynia are more like those of .C. molesta with lanceolate shapes and ventral nerves. Both specimens were collected in a soil type typical of .C. molesta with moist organic silts mixed with sand.

Intermediate Specimens of Carex molesta and Carex festucacea

An itermediate specimen of _c. molesta and .C. festucacea was found at BSM. .C. molesta traits for M:906 include circular to oval perigynia with tapering beaks. ..C. festucacea traits include high inflorescence length; low inflorescence width/length; moniliform spike arrangement (approximate spike tips); spike bases clavate; high number of male cycles (9) on terminal spike; high ratio: terminal spike male cycles/female cycle; no ventral nerves on perigynia.

The specimen labeled F:lOla was collected with M:lOlb at the base of a sand slope (BSM). It was initially analyzed as an unknown and only classified as

.C. festucacea after seeing results of canonical analysis. It most resembles _c. brevi or in overall appearance. Character states for F: lOla normally attributed to

.C. festucacea include narrow achene width (1.0); low ratio: achene 101 length/perigynia length; low perigynium length to widest point/perigynia length

( = 0.44 ); long beak length = 1.42 mm. However, character states normally attribruted to .C. molesta were globose spikes, short inflorescence, high inflorescence width/length, perigynia wider than 2mm, lowest value of female cycles on terminal spike (9, next value 12); lowest lateral female cycles; ovate perigynia bodies (all .C. festucacea specimens had oval bodies); tapered beaks; low scale width/perigynia width ( = 0.48); low scale length/perigynia length ( =

0.79); ventral nerves at base only; and ascending beaks.

Intermediate specimens of Carex brevior and Carex festucacea

Mixed character suites for F:945 include lanceolate to narrowly ovate perigynia with no ventral nerves (perigynia extremely long for .C. festucacea); moniliform spike arrangement; strongly clavate spike bases; spikes oblong, spike beaks ascending to spreading; long achene length and width yet achene is elliptically shaped; the highest lateral male cycles/ female tiers ratio of any specimen of .C. brevior and _c. festucacea; and the lowest scale width/perigynia width for festucacea ( = 0.42 next highest 0.48).

B:104 was collected in a small depression on a large sand dune at BSM.

Many of the characters were within the normal distributions for .C. brevior: moniliform spike arrangement and perigynia ascending to spreading. However, features of the perigynium were more like .C. festucacea: low perigynium length, strongly 3-nerved ventrally low perigynia length, perigynia width under 2 mm, narrow achene width, elliptically shaped achene, high number of dorsal nerves, 3 ventral nerves, low scale length and width, extremely low beak length, high 102 achene length to perigynia length ratio, and the length-to-widest-point-of­ perigynia is near the mean for .C. festucacea. Intermediate specimens of Carex tenera and Carex normalis. At SRCP, I randomly collected a specimen (T:55) growing in the meadow at the edge of an aspen thicket. This specimen had perigynia very similar with those of .C. normalis, collected about 2 meters away growing in gray-loam soil beneath the aspens. The vegetative features were like those of .C. tenera which grew in moist peat or muck soil. 103

CHAPTER 4

DISCUSSION

This study demonstrates that: 1) Carex breyior• .C. festucacea, _c. molesta,

_c. normalis, .,C. tenera. and .,C. tenera var. echinodes are morphologically distinct taxa; 2) there are suites of taxonomic characters useful in classifying and recognizing taxa; and 3) while taxonomic characters vary, most of this variation occurs within populations in predictable patterns.

Taxonomic Classifications

This study was conducted under the premise that current taxonomic classifications for the .,C. brevior group were valid. I based this determination upon the examination of an extensive number of specimens collected throughout the eastern and central United States. I concentrated mainly on the study of historical specimens cited or annotated by authors in their treatments of the _c. straminea aggregate. I was able to use multivariate statistics, specifically

Discriminant Function Analysis (DFA), to test the hypothesis that morphological forms within the .C. brevior group are significantly different. This procedure differs from most taxonomic studies which assume no apriori classification and use Principle Component Analysis to formulate a classification scheme. Random sampling removed any bias in the selection of specimens and enabled me to estimate frequencies of occurrance for morphological forms within populations, between populations, and among taxa. It was only by randomly collecting up to

30 samples per population, from multiple populations, that I was able to determine if morphological differences between taxa were significant. Normal 104

distributions for characters states indicate that the sample size was sufficient to

include most of the variation expressed by the taxa. The results of this study

show that: 1) Iowa representatives of the .C. breyior group are significantly

different from one another, 2) there are suites of correlated morphological

characters capable of distinguishing taxa, and 3) taxonomic characters vary in

predictable patterns according to current theories of Carex morphological

development and cytogenetics.

This study has two limitations: 1) three taxa were collected from only one

or two populations, and 2) only Iowa specimens of the .C. breyior group were

sampled. However, results from taxa with fewer sample sites were in agreement

with results obtained from more extensive collections of .C. breyior and .C.

molesta. In addition, my examination of herbarium specimens indicates that the

Iowa specimens documented in this study accurately represent morphological

forms found throughout the entire geographic distribution of the taxa, i.e., i characters of specimens collected from other regions agree with the character distribution parameters provided in Table 6 for Iowa collections. For example,

~,I Rothrock (1991) has shown Carex festucacea to be a distinct species using

Principal Component Analysis of selected herbarium specimens. Multivariate

analysis of a single population of .C. festucacea in Iowa (BSM) identified

taxonomic characters similar to those selected by Rothrock (1991). Character

means were similar in both studies (within one standard deviation) for

inflorescence length, number of spikes per inflorescence, spike length, perigynium

width, perigynium length from base to widest point, number of ventral nerves,

achene body width/length ratio, and achene width. 105

Taxonomic Characters

This study documents that morphological characters of the .C. brevior group vary within populations in a characteristic pattern. Much of this variation can be explained using current theories of morphological development and cytogenetic evolution. It is likely, based on genetic research of other Carex populations, that most of the variation documented in this study can be attributed to 1) genetic differences between taxa, 2) phenotypic plasticity, and 3) hybridization. These conclusions are probably applicable to the rest of the .C.

straminea aggregate and perhaps the section Ovales as well.

DFA selected taxonomic characters based on statistical measures of significance (Wilk's lambda and the F-values). Yet univariate analysis

demonstrates that these characters overlap among taxa. Therefore, individual

characters cannot distinguish the taxa alone, but must be used in combination with other characters to create character suites cabable of separating taxa. I will

now discuss the taxonomic applications and limitations of these characters.

Yeietatiye Characteristics

Mackenzie (1931-1935) used vegetative characters to create subsectional

divisions within the genus Carex. Subsections Tribuloideae and Festucaceae were distinguished by features of the culm and leaves (number, width, length,

cross-section, sheaths etc.). These vegetative characters are correlated with

taxonotpic features of the perigynium. One exception is .C. normalis which has vegetative qualities of _c. tribuloides but perigynium features similar to .C.

festucacea (representative species of Mackenzie's subsections). Leaf 106 characteristics distinguish the .C. brevior group and subsection Festucacea from subsection Iribuloideae, which includes .C. tribuloides. .C. cristatella. .C. projecta, and .C. muskin~mensis. The Festucaceae have narrower (.s. 4 mm), v-shaped leaves, whose bases are on lower third of the fertile culm. While the

Tribuloideae, usually have wide ( > 5 mm), w-shaped or phlanged shaped leaves, with leaf bases mostly two-thirds the length of the fertile culm. .C. normalis is unusual for the Festucaceae. because it has wide, phlanged shaped leaves, whose bases are up to one-half the culm's length. The perigynia of .C. normalis are characteristic of the Festycacea with ovate perigynia (instead of obovate) which are uniformly winged to the base (instead of wings constricted at mid-point of perigynia, narrowing toward the base).

Some authors make use of color, leaf shapes, leaf ligules and sheaths, as well as the presence and absence of vegetative culms. I found these features useful in the field for comparing one specimen to another, but could not make use of them otherwise. Within a group of morphologically similar taxa such as the .C. breyior group, I have observed that many vegetative characters are unreliable and nearly impossible to quantify for statistical analysis. However,

Rothrock (1991) has quantified several features of the leaf sheath and culm for distinguishing .C. festycacea. .C. albolutescens and .C . .lmliii- Vegetative cu1ms express useful characteristics and should always be examined. However, they are not capable of distinguishing taxa within the

_c. breyior group. When characters of vegetative cu1ms were included in stepwise discriminant analyses they had relatively low discriminating power. For example, 107 the number of leaves on vegetative culms for _c. molesta varied from 3-13 which included the entire range of all the other taxa combined. The width of the largest vegetative culms were smaller than the width of fertile culms for all taxa.

Vegetative culms varied considerably within the genet, while fertile culm widths were consistent. Vegetative features such as ventral sheath surfaces (hyaline vs. green-striated), ligules, leaf cross-sections, etc., are easier to observe on vegetative culms than on senescent fertile culms.

variation of ve&etatiye characters. Size, shape and color of both types of culms varied in similar manner in all taxa. Generally, genets of the same taxa were a darker green in shaded areas while those in open sun were a yellowish­ green. Shaded plants and plants in dense herbaceous cover often had thinner, rounder and more elongated culrns than plants in open and disturbed habitats.

Hyaline portions of ventral sheaths cannot be seen on smaller culrns even from those taxa where this is a characteristic feature. _c. suberecta is described as green-striated to the mouth of the ventral sheaths and this is a consistent feature.

However, I have only seen specimens collected from highly competitive sedge­ meadows bordering fens where the culms are less than 3rnm wide. Typically, .C. tenera and _c. scnparia have hyaline ventral sheaths, but these are often not expressed. The absence of hyaline sheaths can be attributed to poorly developed culms often found in dense herbaceous cover or shaded areas. The dorsal sheath of several taxa, especially _c. breyior and _c. bicknellii, have the green-white mottled color which Mackenzie described as characteristic of _c. normalis. I have found features of the sheath are not reliable enough to be primary key characters in opening leads. 108

Pseudoculms represent one of many morphological stages in the

development of cu1ms in the .C. breyior group. Leaves of these specimens were

clustered at the apex of the culm and surrounded by a sheath originating from the

culm's base. The height of the extended leaves was consistently about 11 em for

all taxa of the .C. breyior group. When the outside sheath was removed and the

leaves were dissected away, I exposed apical meristems on a 3-4 em culm. It

appears that the leaves are clustered within the sheath because the internodes

have not yet elongated. I found vegetative cu1ms at several morphological stages

of development in every population from those still in the coleoptile-like prophyll

to well developed vegetative culms with vascular tissue, nodes and internodes.

True vegetative culms with nodes and internodes were found on nearly every

genet. These were occasionally more numerous than culms with fertile

inflorescences. Vegetative cu1ms are capable of overwintering above ground,

ultimately developing into inflorescences the following spring.

Inflorescence Characteristics

Most of the evidence for the taxonomic classification of Carex is based on

the features of the inflorescence, spike and perigynium. While each taxon

exhibits distinctive forms of these structures, they vary considerably in shape, size

and color. Inflorescence characters are most useful in segregating subgroups

within the .C. breyior group. Useful characters include the average length of the

inflorescence, inflorescence width/length ratio, and the arrangement and number

of spikes.

.11 109

Variation within the inflorescence. Inflorescence morphology is determined primarily by internodal distances. The manner by which spikes are arranged along the rachis of the inflorescence varied in all populations of each taxon from aggregate to moniliform. Morphological extremes were seen and measured in ramets of the same genet. Strongly moniliform spike arrangements in .C. brevior and .C. festugcea were moderately correlated with longer inflorescence lengths. Inflorescence lengths in both taxa were also moderately · correlated with an increased number of male floret cycles on lateral spikes. The number of male floret cycles was correlated with clavate spike bases. I observed internode distances of the axis between male florets with varying lengths. This added to the clavate appearance of spike bases. These variations suggest that inflorescence morphology is controlled by growth hormones which affect the internode lengths of inflorescences and spikes (Smith 1967 1969).

Spike Characteristics

Spike shape and size are the most useful taxonomic characteristics of the inflorescence. Spike appearance is influenced by the number of florets, the internodal distances between florets, and the ratio of male to female florets on the spike. Spreading perigynium beaks give the spike an echinate or bristly appearance with round apices and bases. Ascending beaks give the appearance of acute spike tips and elliptic spikes. Spike length and lengths of the male and female portion of the spike are useful but highly variable characters and must be used cautiously. The number of male florets and their internode length determine whether a spike base is round or clavate. 110

variation of spike characters. Florets develop spirally around the rachilla.

The spiral ascends from right to left as viewed laterally--clockwise if viewed from

the top. The spikes are indeterminate. Empty pistillate scales are always present

at the apex because the perigynia never develop. The number of female cycles

varies in all taxa and spike shapes can be attributed to development. An

increasing number of female florets will change the overall shape of the spike

from globose to elliptic (or obovate to oblong, depending on the number of male

florets). Whether perigynia beaks are appressed or spreading may depend on the

number of perigynia per measured length of the rachilla. Shorter internode

lengths between florets compresses perigynia on spike axes causing their beaks to

spread apart as the perigynia mature. .,C. breyior has longer spike axes, relatively

few perigynia, and ascending beaks. In contrast, ..C. molesta has a greater number

of perigynia compressed on shorter spike axes causing the perigynia to spread

widely giving the spike a bristly or echinite appearance. Long perigynium beaks,

short spike lengths and proportionally low numbers of male florets accentuates

the round appearance of spikes in ..C. normalis, .,C. molesta and ..C. tenera var.

echinodes. \·.• :! ! Meristems of the inflorescence develop into male florets, female florets or '1 •j elongate into spikes (Smith 1966). I found sexual structures in many specimens l :l where they do not normally occur. In all taxa, I frequently observed stamens in I positions where perigynia or spikes normally developed. Also, isolated male II florets were found in various positions between the second and seventh cycles of ~ the female portion of the spike. At the base of each spike, there is a subtending - 111

bract on the rachis. The bract of the lowest spike is usually setaceous and often

exceeds the inflorescence. Stamens are found in nearly every spike bract except

the lowest. Occasionally the spike axis aborts and no spike develops leaving an

empty bract. More frequently, additional spikes form in the subtending bract of

the terminal spike. These extra spikes have one to several perigynia. One such

spike developed from a male floret position in the second cycle of the terminal

spike (F:893). These observations support morphological evidence which

indicates that Carex florets are actually reduced spikelets with an aborted rachilla.

Smith (1967) experimented with the addition of plant hormones to developing

meristems. He determined that various hormones could alter the outcome of the

meristems and affect the numbers of male florets, female florets, or lateral spikes.

Club-shaped or clavate spike bases are a diagnostic feature for

.C. festucacea. However, spike bases vary considerably in all taxa and this distinctive feature is not always present in .C. festucacea. The spike morphology

is determined by the number of floret cycles around the axis and internode

distances. The number of cycles of staminate florets varies in all taxa, especially

in terminal spike bases. However, the number of male cycles in lateral spikes is

more consistent Therefore, lateral spike bases exhibit more useful

characteristics.

Scale and Bract Characteristics The size of the scale in relation to the perigynium is a distinctive quality.

The ratio: scale width/perigynium width segregates .C. brevior and .C. molesta

from the rest of the taxa, while the ratio: scale length/perigynia length separates 112 this pair from each other. Scale length/perigynium length also distinguishes _c. tenera var. echinodes due to its longer perigynia. I cannot discern any differences in color, nervation or shape of the pistillate scales in any taxa, except _c. brevior, whose pistillate scales are usually longer than other taxa. _c. festucacea was reported by Dewey (1824) to have staminate scales with accuminate tips, but this is not true for Iowa specimens.

Variation of scale and bract characters. Scales vary in shape and size depending on position. Staminate scales near the spike base are shorter with truncate or obtuse tips. Scale length becomes normal with acute tips by the third cycle of male florets. Bracts subtending spikes are mostly indistinguishable from staminate scales except for the lowest bract which is usually setaceous. Often the bristled tip of the lowest spike bract equals the length of the inflorescence.

Peri&YJlium Characteristics

Perigynium characteristics account for 39% of the total number of characters analyzed in this study. Stepwise Discriminate Analysis of the entire dataset selected a total of 37 characters with discriminant powers; about 35% of these were perigynium traits. Historically, all specific and varietal classifications in Carex are based on the perigynium. Therefore, using traditional species concepts, distinguishing characteristics of a taxon's life history, habitat, rhizome, culm, leaf or inflorescence must be correlated with recognizable features of this unique structure. Critical examination of perigynia is necessary for accurate identifications of specimens in the _c. brevior group. But the perigynium can never be the only criteria for recognition because its distinguishing features vary 113 considerably within populations, or even within genets, or they may be absent.

The most characteristic perigynia are found between one-half to two-thirds the spike length as measured from the base.

The size and shape of the perigynia account for much of the variablity between taxa. Average perigynium lengths of the .C. breyior group range from 3 trim to 4.25 mm. However, perigynia lengths vary even within the same spike and their ranges of variation overlap among the taxa. Perigynium width separates the

.C. breyior group into two subgroups. Taxa with lanceolate or narrowly ovate perigynia seldom exceed 2 mm in width, i.e., _c. breyior and .C. molesta with circular or elliptic perigynium shapes characteristically have perigynia widths 2 mm or greater. Similarly, the location of the widest point on the body of the perigynium is also a good taxonomic character. The ratio of the length-to-the­ widest-point divided by the perigynium length is a reliable measure of this characteristic shape. Perigynium shape separates .C. tenera var. echinodes and .C. nonnalis. which are narrowly ovate, from .C. breyior. whose perigynia are consistently circular. .C. molesta and .c. festucacea tend to have elliptical perigynium bodies while .C. tenera perigynia tend to be ovate. Perigynium shapes were also measured qualitatively using an adaptation of standard shapes provided by Radford et al. (1974). The two methods for enumerating perigynium shapes essentially double-weighted the discriminating power of this feature. However, removal of the second qualitative measure did not significantly alter the canonical distances between taxon classes, while its inclusion gives greater clarity in defining shape characteristics. 114

The size and shape of perigynium beaks are good taxonomic characters.

In order to reduce ambiguity, I measured beak length from the apex of the achene to the tip of the perigynium teeth. Within taxa, beak length is directly proportional to perigynium length and· beak lengths overlap among taxa.

Therefore, it is not a distinctive trait. A more useful taxonomic character is the ratio: beak length/perigynium length. This feature separates .C. breyior and .C.

festucacea from other taxa. Beak shape is based on the outline of the perigynium

from the mid-point to the beak apex. Tapered beaks are characteristc of

.C. tenera. _c. tenera var. echinodes. .C. molesta. and .C. normalis. Constricted beaks curve inwards above the achene, forming short, narrow beaks charcteristic

of .C. breyior and .C. festucacea. Beak shape is a good secondary character.

The quality and number of perigynium nerves over the achene have

taxonomic merit. However, nerves are difficult to see on immature perigynia.

Strongly-nerved perigynia have relatively thick nerves elevated above the surface of the perigynium. Fine nerves are imbedded within the tissue of the perigynium

or just slightly raised. Nerve quality is subjective, therefore comparisons must be

made with known specimens. .C. breyior typically does not express perigynium

nerves on the ventral surface and is the only taxon of the group readily identified

by nerve counts.

Characteristics of the Achene

Achene width and achene shape are useful for separating .C. brevior and

.C. molesta from other taxa. like other characters, they have separate means with

overlapping ranges of variation. Ratios of achene width/perigynium width and

achene length/perigynium length are good secondary characters. 115

Sources of Mor.pholo&ical Variation

Most of the morphological variation expressed by Iowa specimens of the .C. breyior group is found at the population level because there are no detectable differences in population patterns within taxon clusters. This is indicated by the uniform distribution of population codes within taxon clusters in canonical plots

(Figures 9-11) and normal distributions of single .characters (Figure 5). Also, specimens of sympatric populations of ,C. brevior and ,C. molesta occupied somewhat intermediate positions in canonical plots between allopatric populations within the taxon clusters of these two species (Figure 12 and Table 15). I believe the variation documented in this study may be attributed to phenotypic plasticity and hybridization.

Phenotn>ic Plasticity

Studies of microspecies within species aggregates have shown that there is low genetic variation in Carex populations (Whitkus 1988; Bruederle and

Fairbrothers 1986; Watetway 1990; Bruederle and Jensen 1991). Whikus (1988) demonstrated that microspecies reproduced primarily by selfing resulting in homozygous populations with low genetic diversity. H populations of the ,C. breyior group exhibit low genetic diversity as has been documented for other species aggregates, then the variation recorded within populations is probably due to phenotypic plasticity controlled by environmental gradients. These conclusions are supported by studies of developmental morphology in the genus Carex (Smith

1966 1967 1969). Based on the results of this study and past research, I have concluded the variation found within Iowa taxa is largely due to phenotypic 116 plasticity, while variation among the taxa is due to genetic differences probably maintained by pre-zygotic reproductive barriers and different habitat preferences.

Hybridization

An additional and substantially different type of morphological variation is observed at sympatric population sites. Specimens from sympatric populations occupy intermediate positions between taxon clusters on canonical plots. In addition, a disproportionate number of specimens from sympatric sites are found in hemispheric regions of taxon clusters which are closest to the sympatric taxon.

Removal of specimens collected at sympatric sites would eliminate overlap of clusters, increase the distance between cluster centroids and result in higher F­ values of significance. Canonical plots demonstrate that specimens from each population are homogeneously distributed throughout each cluster, i.e. no secondary patterns are evident within taxon clusters. However, allopatric populations (single species sites) for .C. breyior and .C. molesta form a tighter pattern around cluster centroids while specimens from sympatric populations

(multiple species sites) tend to form looser patterns in peripheral regions intermediate between taxon clusters (Figure 12). Nearly all the specimens plotted within this intermediate zone were specimens collected in close proximity to members of another taxa {Table 15). While suites of correlated characters separate taxa, intermediate specimens from sympatric sites have mixed character suites, i.e., they express character states attributed to both taxa found at the site.

These are difficult specimens to classify. 117

The canonical pattern of .C. festucacea specimens, collected at BSM, is characteristic of a sympatric population. There is a tight grouping of specimens near the taxon centroid, and a second more diffuse grouping in a narrow peripheral segment forming an intermediate band between associate species, _c. brevior and .C. molesta.

The presence of mixed character suites within specimens collected at sympatric sites may be evidence of hybridization or introgression. Cytogenetic research has shown that hybridization between morphologically similar species with small differences in chromosome number is common in Carex. Tanaka

(1940 1949) suggests naturally occurring hybridization may be a common form of speciation within Carex. The frequent occurance of specimens with mixed character states suggests that hybridization is occurring between taxa of the Carex breyior group.

Although inconclusive, evidence from field observations of .C. tenera var. echinodes combined with morphological analysis supports the conclusion that this taxon may have originated from a naturally occurring hybrid cross between .C. tenera and .C. normalis. .C. tenera var. echinodes displays a mixed combination of character states with perigynia very similar to .C. nonnalis, but vegetative characters similar to .C. tenera. .C. tenera was found in moist high-organic soils formed under prairie vegetation, while .C. normalis was found in well-drained, grayish-loam soils formed under prairie and hardwood forest. A population of _c. tenera var. echjnodes was collected at a shaded hill-side seep in Martin County

Park. This habitat is unique to .C. tenera var. echinodes. Specimens of .C. tenera 118 var. ecbinodes are plotted in between .C. tenera and .C. normalis on the second canonical axis (Figures 6 and 7). A specimen of .C. tenera (T:55) was collected 3 meters away from two specimens of .C. normalis at SRCP. T:SS closely resembles .C. tenera var. echinodes. It was found growing in an intermediate or transition zone between peat soil and upland loam where .C. tenera and .C. normalis were found. 119

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Tanaka, N. 1949. Chromosome studies in the Genus Carex. with special reference to aneuploidy and polyploidy. Cytologia 15: 15-29.

Torrey, J. 1836. Monograph of North American Cyperaceae. Ann. Lyceum Nat. Hist. New York 3: 239-246, 394-396.

Townsend, F. 1885. Homology of the. floral envelopes in Graminae and Cyperaceae. Jour. Bot. 23: 65-74.

Tuckerman, E. 1843. Enumeratio methodica Caricum quarundam: species recensuit et secundum habitum pro viribus disponere tentavit. Printed by author (Gray 1843), 21 pp.

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Wahl, H. A 1940. Chromosome numbers and meiosis in the genus Carex. Amer. J. Bot 27: 458-470.

Wahleilberg, G. [1803] 1806. A monograph of the genus CVex. Ann. Bot. 2: 112-144. Translated from Kong. Vetenskaps Academiens nya Handlingar. 24: 138-170. 126

Waterway, M. J. 1990. Genetic differentiation and hybridization between Carex &fDOdynama and ,C. mendocinensis (Cyperaceae) in California. Amer. J. Bot. 77: 826-838.

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V XIGN3ddV

LZI 128

CAREX STRAMINEA Willdenow ex Schkuhr Willdenow (1805)

.C. straminea Schkuhr ex Willdenow Schkuhr (1801) No. 49, Plate G, Fig. 34 . .C. straminea Schkuhr ex Willdenow Wahlenberg (1803) No. 38, p 145 [p 119 of (1806) translation] ( = C. brevior?) .

.C. straminea Schkuhr ex Willdenow Willdenow (1805) No. 73, p 242 .

.C. straminea Schkuhr ex Willdenow Schkuhr (1806) No. 62, Tab. Xxx, Fig. 174.

_c. straminea Schkuhr ex Willdenow Muhlenberg (1817) p 229. .c. 5traminea Schkuhr ex Willdenow Schweinitz (1824) No. 35 ( = .C. brevior?).

.c. stramin~a Willdenow ex Schkuhr Schweinitz (1826) No. 34 ( = .C. brevi or?). _c. straminea Wahlenberg non Willdenow Dewey (1826a) p 157. _c. straminea Willdenow ex Schkuhr Torrey (1836) No. 37, p 395 ( = .C. brevior).

Vi~nea straminea Rafinesque (1840) [The Good Book and the Amenities of Nature p 27: See Merrill 1949] .

.C. straminea Willdenow ex Schkuhr Tuckerman (1843)

.C. straminea Willdenow ex Schkuhr Carey (1856) No. 41, p 516 (=_c. brevior).

_c. straminea var. aperta F. Boott Boott (1862) p 120, Fig. 385 (p. p.) .

.C. straminea var. aperta [forma] major Olney Olney (1870) Exsiccatae Carices Americanae-Boreali (p. p.) .

.C. straminea var. aperta F. Boott Bailey (1886) p 152 (p. p.) . .C. straminea var. aperta F. Boott Bailey (1889 1890) p 24. and p 622 (p. p.).

~ tenera Dewey Mackenzie (1896) No. 200, Fig. 870, p 358 .

.C. tenera var. Richii Fernald Fernald (1902) No. 12, p 475, Figs. 33 and 34. 129

_c. hormathodes var. Richii Fernald Fernald (1906 1908) pp 165-166, and No. 12, p 220, Fig. 358 .

.C. straminea var. tenera forma Richii Kukenthal Kukenthal (1909) No. 167, p 206 .

.C. hormathodes Fernald Mackenzie (1913) No. 71, p 384, Fig. 938.

_c. Richii Mackenzie Mackenzie (1931 1940) No. 180, p 160, and plate 180.

_c. straminea Willdenow ex Schkuhr Fernald (1950) No. 94, p 327, Fig. 597.

CAREX FESTUCACEA Schkuhr ex Willdenow (1805)

_c. festucacea Schkuhr ex Willdenow Willdenow (1805) No. 72, p 242.

_c. festucacea Schkuhr ex Willdenow Schkuhr (1806) No. 62, p 23, Fig. 173.

_c. festucacea Schkuhr ex Willdenow Muhlenberg (1817) p 249.

_c. festucacea Schkuhr ex Willdenow Schweinitz (1824) p 66.

_c. festucacea Schkuhr ex Willdenow Dewey (1824) No. 15, p 96.

_c. festucacea Schkuhr ex Willdenow Schweinitz (1826) No. 37, p 316.

_c. festucacea Schkuhr ex Willdenow Torrey (1836) No. 36, p 394.

_c. straminea var. festucacea Gay (1838) [Ann. Sci. Nat. (II) x. 363 [Bailey (1889) footnotes that description founded on ..C. normalis and ..C. adusta at Kew] .

.C. straminea var. festucacea Gay Tuckerman (1843)

_c. festucacea Schkuhr ex Willdenow Carey (1856) No. 39, p 516.

_c. straminea var. festucacea Gay Boott (1862) p 120, Fig. 386.

_c. straminea var. festucacea Gay Carey (1867) No. 44, p 580.

_c. straminea var. (No. 1) Bock (1875) [Linneae 39: 117].

_c. straminea Willdenow ex Schkuhr Bailey (1886) [p. p.] No. 283, p 149.

_c. straminea var. brevior Dewey Bailey (1889) [p. p.] No. 31, p 22. 130

..C. straminea var. brevior Dewey Bailey (1890) [p. p.] No. 132, p 622.

Not ..C. festucacea Willdenow sensu Mackenzie (1896) No. 201, p 359, Fig. 871. (=..C. brevior and ..C. merritt-fernaldii)

..C. straminea Willdenow Mackenzie (1896) [p. p.] No. 198, p 358, Fig. 868 .

..C. straminea Willdenow Fernald (1902) [p. p.] No. 11, p 474.

Not _c. festucacea Schkuhr sensu Fernald (1902) No. 16, p 477, Figs. 47, 48. ( = ..C. merritt-fernaldii) .

..C. straminea Willdenow Fernald (1908) [p. p.] No. 11, p 219.

Not C. festucacea Sckuhuhr sensu Fernald (1908) No. 17, p 220-221, Fig. 363 ( = _c. merritt-fernaldii) .

..C. straminea var. festucacea (Schkuhr) Tuckerman Kukenthal (1909) p 206 .

..C. straminea Willdenow Mackenzie (1913) [p. p.] No. 66, p 382.

Not _c. festucacea Schkuhr sensu Mackenzie (1913) No. 69, p 383, Fig. 936 ( = ..C. brevior and ..C. merritt-fernaldii) .

..C. festucacea Schkuhr ex Willdenow Mackenzie (1915) p 608 .

..C. festucacea Schkuhr ex Willdenow Mackenzie (1931) No. 165, p 150.

_c. festucacea Schkuhr ex Willdenow Mackenzie (1940) No. 165 .

..C. festucacea Sckhuhr ex Willdenow Fernald (1950) No. 81, p 325, Fig. 583.

_c. festucacea Schkuhr ex Willdenow Rothrock (1991) No.2, p 63, Figs. 2-5.

CAREX TENERA Dewey (1824 & 1825)

..C. tenera (Mihi) Dewey (1824) No. 17, p 97 and Dewey (1825) Plate III, Fig. 9.

_c. straminea var. minor Dewey Dewey (1826) p 318, Tab. N, Fig. 45 .

..C. tenera Dewey Schweinitz (1826) No. 41, p 319 .

..C. straminea var. minor Dewey. Torrey (1836) No. 38, p 395 .

..C. straminea var. minor F. Boott ex Hook (1839) [Flora Boreal America 2: 131

215: See Gray Herbarium Index] .

..C. straminea var. tenera (Dewey) Barratt (1840) [N. Amer. Carie. no. 51: see Gray Herbarium Index].

Diemisia tenera Rafinesque (1840) [The Good Book and the Amenities of Nature: see Merrill (1949)] .

..C. straminea var. moniliformis Tuckerman (1843) [p. p.] .

..C. festucacea var. tenera Carey (1856) No. 39, p 516 [Also 1st ed. Gray's Manual (1848) p 545] (=_c. hormathodes?) .

..C. straminea var. tenera Boott (1862) p 120, Fig 384.

_c. mirabilis var. tenera (Dewey) L. Provancher (1863) [Flora Canada 2: 648].

_c. tenera (Dewey) forma erecta Olney (1870) Exsiccattae fasc. ii, No. 14 . ..C. straminea var. (No. 2) Boeckl. (1875) [Linneae 39: 117]. _c. straminea Willd Bailey (1886) [p. p.] No. 283, p 149. _c. straminea Willdenow Bailey (1889) No. 31, p 21-22. _c. straminea Willdenow Bailey (1890) No. 132, p 621.

..C. moniliformis (Tuckerman) Britton (1890) [Cat. plants N.J. p 278].

_c. straminea Willdenow Mackenzie (1896) No. 198, p 358, Fig. 868. _c. straminea Willdenow Fernald (1902) No. 11, p 474, Fig. 28 and 29 .

..C. straminea Willdenow Fernald (1908) No. 11, p 219, Fig. 354.

_c. straminea Willdenow Kukenthal (1909) No. 167, p 205, and p 204, Fig 34, E & F.

_c. straminea Willdenow Mackenzie (1913) No. 66, p 382, Fig. 932. _c. tenera Dewey Mackenzie (1915) p 606-607. _c. tenera Dewey Mackenzie (1931) No. 162, p 148.

_c. tenera Dewey Mackenzie (1940) No. 162 (perigynium illustration is ofvar. 132

echinodes not of Dewey's lectotype).

_c. tenera Dewey Fernald (1950) No.80, p 325, Fig. 581.

CAREX TENERA (Dewey) V AR. ECHINODES (Fernald) Wiegand Fernald (1902).

_c. straminea var. echinodes Fernald (1902) No. 11, p 474, Fig. 30.

_c. straminea var. echinodes Fernald (1908) No. 11, p 219, Fig. 355

_c. straminea forma echinodes Fernald Kukenthal (1909) No. 167, p 206.

_c. festucacea var. echinodes (Fernald) Farwell (1923) [Papers Mich. Acad. 2: 17]

.C. tenera (Dewey) var. echinodes (Fernald) Wiegand (1924) [Rhodora 26: 2.].

_c. tenera Dewey Mackenzie (1931) [included in description] No. 162, p 148.

_c. tenera Dewey Mackenzie (1940) No. 162 [illustration of perigynium is of var. echinodes].

_c. tenera var. echinodes Fernald Wiegand Fernald (1950) No. 80, p 325, Fig. 582.

CAREX BREVIOR (Dewey) Mackenzie (1915) .

.C. straminea Willdenow Wahlenberg (1803)? No. 38, p 145 [p 119 of 1806 translation].

_c. straminea var. brevior Dewey (1826) p 158.

_c. straminea Willdenow Torrey (1836) No. 38, p 395.

_c. straminea var. Schkuhrii Gay (1838) [Ann. Sci. Nat. 10: 364] .

.C. straminea var. intermedia Gay (1838) [Ann. Sci. Nat. 10: 363. Bailey (1889) footnotes that Kew Herbarium has C. brevior and _c. silicea. However, Gray Herbarium Index applies it to .C. festucacea].

_c. straminea var. Schkuhrii (Mihi) Tuckerman (1843) [This is cited as the typical form for _c. straminei!, but should be applied to C. brevior].

_c. straminea Willdenow Carey (1856) No. 41, p 516. _c. straminea var. typica Boott (1862) p 121, Fig. 387. 133

_c. straminea Willdenow Bailey (1886) [p. p.] No. 283, p 149.

_c. straminea var. brevior Dewey Bailey (1889) No. 31, p 22.

_c. straminea var. brevior Dewey Bailey (1890) No. 132, p 623.

_c. festucacea Schkuhr Mackenzie (1896) [p. p.] No. 201, p 359, Fig. 871 (illustration of _c. merritt-fernaldii).

_c. festucacea var. brevior (Dewey) Fernald (1902) No. 16, p 477, Figs. 49-51.

_c. festucacea var. brevior (Dewey) Fernald (1908) No. 17, p 221, Fig. 364.

_c. straminea var. brevior Dewey Kukenthal (1909) No. 167, p 207.

_c. festucacea Schkuhr sensu Mackenzie (1913) No. 69, p 383, Fig. 936.

_c. brevior (Dewey) Mackenzie comb. nov. Mackenzie (1915) p 603-605.

_c. brevior (Dewey) Mackenzie ex Lunell (1915) [American Midland Naturalist 4: 235 - Not listed in text!].

_c. brevior (Dewey) Mackenzie (1931) No. 167, No. 151.

_c. brevior (Dewey) Mackenzie (1940) No. 167.

_c. brevior (Dewey) Mackenzie Fernald (1950) No. 86, p 326, Fig. 588.

C. NORMALIS (Dewey) Mackenzie Dewey (1936)

_c. mirabilis Dewey (1836) p 63, Tab Bb. Fig. 92. [Not C. mirabilis Host (1809)].

_c. straminea var. mirabilis (Dewey) Tuckerman (1843).

_c. festucacea var. mirabilis (Dewey) Carey (1856) No. 59, p 516 [Also 1st ed Gray's Manual (1848) p 545].

_c. cristata Schw. Kunze (1851) [p. p.] [Supplement to the Reidgraser] [pl. 44, Fig. a, e, f.

_c. cristata Schweinitz Boott (1862) [p. p.] No. 276. p 117, Fig. 374.

_c. cristata var. mirabilis (Dewey) Carey? ex Gray (1867) No. 41, p 580. _c. la~opodioides var. mirabilis Olney (1870) Exsiccattae fasc. ii no. 9. 134

.C. tribuloides var. cristata Bailey (1883) [See Mackenzie (1931)] .

.C. straminea var. mirabilis (Dewey) Tuckerman Bailey (1886) ["mostly, some specimens of Dewey and others listed under .C. tribuloides var. cristata"] No. 283, p 150 .

.C. mirabilis Dewey [and including] var. perlon&a Fernald Fernald (1902) No. 10, p 473, Figs. 25-27 .

.C. mirabilis Dewey [including] var. perlon~a Fernald Fernald (1908) No. 10, p 219, Figs. 352-353 .

.C. straminea var. mirabilis Tuckerman [including] forma perlon~a Fernald Kukenthal (1909) No. 167, p 207 .

.C. normalis (Dewey) Mackenzie (1910) p 244 .

.C. normalis var. perlonga (Fernald) Burnham (1919) [Torreya 19: 131] .

.C. normalis (Dewey) Mackenzie (1931) No. 164, p 149 .

.C. normalis (Dewey) Mackenzie (1940) No. 164 .

.C. normalis forma perlonga Fernald (1942) p 285 .

.C. normalis (Dewey) Mackenzie [including] forma perlon~a Fernald (1950) No. 79, p 324-325, Figs. 579-580.

C. molesta Mackenzie ex Bright (1930) .

.C. molesta Mackenzie ex Bright (1930) p 20 [See Rothrock (1978)] .

.C. molesta Mackenzie (1931) No. 166, p 151.

_c. molesta Mackenzie (1940) No. 166 .

..C. brevior var. molesta (Mackenzie) F.C. Gates (1940) p 135 .

.C. molesta Mackenzie Fernald (1950) No. 87, p 326, Fig. 589 .

..C. brevior (Dewey) Mackenzie Gleason and Cronquist (1952) [p. p.] No. 80, p 325 [authors consider the taxon a possible hybrid .C. brevi or x C. normalis].